Lynn Margulis and Dorion Sagan

Foreu-ord b), Niles Eldredge

I
WHAT IS LIFE?

e7
.G
t
The publisher gratefully acknowledges the generous
contribution to this book provided by the following
organizations and individuals:

Elizabeth Durein
Orville and Ellina Colub
David B. Cold Foundation
Moore Family Foundation

and by the Ceneral Endowment Fund of the

Associates of the University of California Press.
LYNN MARCULIS
DORION SACAN

FOREWORD BY

NlLES ELDREDCE

A Peter N. Nevraumont Book

UNIVERSITY OF CALIFORNIA PRESS

Berkeley and Los Angeles

 University of California Pres
Berkeley and Los Angeles, California

Published by arrangement with Simon & Schuster.

First paperback printing zooo.

@ 1995 by Lynn Margulis and Dorion Sagan

Foreword O 1995 by Niles Eldredge
Glossary @ zooo by Lynn Margulis and Dorion Sagan

The Sources of Illustrations ortpage 269 is regarded as

an extension of the copyright page.

Library of Congress Cataloging-in-Publication Data

Margulis, Lynn, r93 8-

What is life? / Lynn Margulis and Dorion Sagan ;

foreword by Niles Eldredge.

'A Peter N. iv"rrrr-or,PUootIi

Originally published: NewYork :
Simon & Schuster, @ rqgS. With new glossary.
Includes bibliographical reGrences (p. ) and index.
ISBN o-5zo-zzozr-8 (pbk. : alk. paper)
I. Life @iology) z. Biology-Philosophy.
3. Biological diversity. 4. Life-Origin.
I. Sagan, Dorion, rg59- II. Tide.
QH5or.M35 zooo
57o-dczr oo-o25833

Manufactured in Canada

09 08 o7 o6 oj 04 03 02 ot oo
ro 9 I I 6 S 4 3 z r

The paper used in this publication meets the

minimum requirements of ANSI/NISO 239.48-t9gz
(R rqSZ) (Permanenrc of Papu). @
 Contents

LIST OF ILLUSTRATIONS vil

FOREWORD
Undreamt Philosophies, by Niles Eldredge xi

LIFE: THE ETERNAL ENIGMA ln the Spirit of Schrodinger ' Life's

Body ' Animism vs. Mechanism ' Janus among the Centaurs ' Blue iewel '
ls There Life on Mars? . Life as Verb . Self-Maintenance ' The Autopoietic
Planet . The Stuff of Life ' Mind in Nature 'l

LOST SOULS Death: The Great Perplexer The Breath of Life

Cartesian License . Entering the Forbidden Realm ' Cosmic Wiggles '
The Meaning of Evolution . Vernadsky's Biosphere ' Lovelock's Gaia 33

ONCE UPON A PLANET Beginnings . Hell on Earth ' Spontaneous

.
Generation ' Origins of Life "stumbling Forward" ' Metabolic Windows '
The RNA Supermolecule . Cells First 57

MASTERS OF THE BIOSPHERE Fear of a Bacterial Planet'Life ls

Bacteria . The Metabolically Gifted ' The Cene Traders ' Our Splendid Kin '
From Plenty to Crisis . Breakfast Ferment' Creen, Red, and Purple Beings '
Oxygen Excitement . Quintessential Polluters, Quintessential Recyclers '
Living Carpets and Crowlng Stones 87

PERMANENT MERGERS The Creat Cell Divide' Five Kinds of Beings '
Twists in the Tree of Life . Squirmers ' Strange New Fruit ' Wallin's
Symbionts' Multicellularity and Programmed Death 'Sexual Genesis in the
Microworld, or When Eating Was Sex 'The Power of Slime 113

THE AMAZING ANIMALS The (Bower) Birds and the (Honey) Bees '
What ls an Animal? ' Great-Crandparent Trichoplax ' Sex and Death '
Cambrian Chauvinism ' Evolutionary Exuberance' Messengers 145
FLESH OF THE EARTH The Underworld . Kissing Molds and Destroy-
ing Angels .
Cross-Kingdom Alliances . Underbelly of the Biosphere .
Hitchhiking Fungi, Counterfeit Flowers, and Aphrodisiacs . Hallucinogenic
Mushrooms and Dionysian Delights .Transmigrators of Matter 171

THE TRANSMUTATION OF SUNLIcHT Green Fire . The Accursed

Share . Ancient Roots . Primeval Trees . Floral Persuasion . Solar Economy
193

SENTIENT SYMPHONY A Double Life . Choice . Little Purposes .

Butler's Blasphemy . Habits and Memory Existence's Celebration
Superhumanity . Expanding Life . Rhythms and Cycles 213

EPILOGUE 245

NOTES 247

GLOSSARY 255

ACKNOWTEDGMENTS 265

SOURCES OF ILLUSTRATIONS 269

INDEX 271
 lllustratio ns

PLATES (following page 1441

ra. Earth in space

rs. Mycoplasma
2. Intestine of beetle larva (Pachnoda)
3. X-ray photograph of the sun
4. Cylinders of phospholipids forming liposomes
5A. Patterns of growth, Proteus mirabilis bacteria
5n. Dissipative structure in a Belousov-Zhaborinsky reaction
6. Multicellular "trees" of Myxococcus
7. Chromatium uinosum, purple sulfirr bacteria
8. Fischerella, cyanobacterium
9. Stromatolites in Shark Bay, Australia
roa and s. Petrified fossil stromatolite compared
to a living microbial mat
tr. Mesodinium rubrum (Myrionecta rubra), a cliate
rz. Telophase in the mitotic cells of Haemanthus sp.,
African blood lily
13e,. Chlamydomonas niualis, snow algae, in Antarctica
r3r. Chloromonas sp., snow algae, and filamentous fungi
t3c. Chlamydomonas niualis, microscopic view
r4. Voluox colonies
r 5.Adult Lima scabra, scallop
16. Embryo of Drosophila melanogaster, fruit fly
r7. Undulipodium cross section

vil
 vilt lllustrations

r 84. Angler fish with bioluminescent tissue

r 8n. Bioluminescent bacteria colonies
19A. Russula paludosa, forest mushroom

ryr. Schizophyllum commune) fungus

zo. A chloroplast
zr. Quaking aspen in San Juan Mountains, Colorado
zz. Diatorn meiosis and gamete formation
23. Glossopteris scutum, fossil seed fern
24. Pollen tubes deliver male nuclei to Gmale egg nuclei
25. Papaver somnferwm, poppy
26. Satellite image of Earth showing major zones of vegetation

FIGURES

r. Erwin Schrcidinger
z. Comparison of atmospheres of Earth, Venus,
and Mars 2'l
3. Fluctuations of carbon dioxide in the northern
hemisphere 24
4. Oxalic acid crystal in a sea squirt 26
5. Emiliana huxleyi, a coccolithophorid 53
6. Three-way genetic exchange among bacteria 95

7. Comparison of prokaryote and eukaryote cells 115

8. Trichonympha. a ptotist '122

9. Spirochetes become undulipodia 124

ro. Stages of mitosis 125
rt. Naegleria, a protist 138
rz. Stephanodiscus, a diatom 143
r3. Sexual life history of an animal 146
14. Eschiniscus blumi, a "water bear" 169
r 5. Sexual life history of a fungus 173
16. Gall on a twig of Quercus, oak tree 180
 lllustrations tx

r7. Sexual life history of a plant 194

r8. Magnetosomes in a magnetotactic bacterium 220

TAB LES

Minerals produced by life 2A-29

Earth history tinrelines 52-AO
 Foreword UN DREAMT PH ILOSOPH IES

Why has evolution crafted a sentient species? Why did our con-
sciousness, our realization of our very existence, evolve? What pur-
pose does it serve? I am persuaded by behaviorist Nicholas Hum-
phries's conjecture that, in being able to consult their inner selves,
our ancestors gained insight on the minds of their mates, offspring,
and other members of their social bands. Klowing thyself is the
best way to knowing others, and thus an advantage in negotiating
the complexities of daily social life.
We humans are, of course, animals. I have long thought that the
very best insight into what it means to be a living, breathing ani-
mal is simply to consider one's very own life. However far our cog-
nitive, cultural capacities have taken us from traditional existence
within local ecosysterN, we nonetheless still obtain energy and food
to develop, grow, and maintain our corporeal existence. Many of
us (perhaps too many of us) also engage in reproduction. As Lynn
Margulis and Dorion Sagan tell ts in What Is LiJe?, the business of
maintaining corporeal existence and reproducing are quintessential
activities, the very hallmarks of life. To know oneself as an organ-
ism, then, is to establish quite a few of the very basics of all living
systems.
But humans, of course, do not constitute the entire biological
universe. 'We are but one species of tens of millions now inhabit-
ing planet Earth. And so we cannot expect to divine all of life's
mysteries, all the different nuances of what it means to be alive,
simply by consulting our inner selves. There are inherent limits to
the revelatory principle of knowing thyself in order to know the
world. But even I, a seasoned practitioner in evolutionary biology,

xt
 Foreword

was not fully prepared for the wild spectrum of life presented to
us by Margulis and Sagan in What ls Life? For in these pages we
meet organisms vastly different from ourselves. And we encounter
ways of thinking about life that could not possibly arise from sim-
ple introspection.
What Is Life? is a feast of biological and intellectual diversity. Here
we meet microbes-microscopic organisms-for which oxygen is
a poison, and others who "breathe" sulfur compounds. And still oth-

ers which feed on hydrogen and carbon dioxide using neither the
-We
energy from sunlight, nor that from the flesh of others. encounter
bacteria routinely exchanging genetic materials with other species-
'We
even after billions of years of evolutionary separation. see the
entire outer rind of Earth portrayed in convincing fashion as a sin-
gle, mega-living system. And we learn that the evolutionary process
that has produced this prodigious array has done so in astonishing
ways-melding separate, simple organisms more than once to pro-
duce more complex descendant species. And therein lies a partic-
ularly interesting saga of intellectual sleuthing and derring-do.
Darwin taught us that all of life is descended from a single com-
mon ancestor.In What k Life? Margulis and Sagan tell us the at71az-
ing fact that not only are our own mammalian, nucleated ("eu-
karyotic") cells descended from ancient bacteria, they are literally
amalgams of several different strains of bacteria. Amazing! Stranger
than fiction! And undreamt of in traditional biological philosophies-
until Lynn Margulis began her research a quarter of a century ago.
Lynn Margulis has achieved what every scientist dreams of, but
few are destined to accomplish: she has rewritten the basic textbooks.
She conceived of a logical, yet audacious explanation of an out-
standing fact. Human cells, like those of all animals, the eucalyptus
tree and the mushroom, have most, but not all, of their DNA cor-
ralled into a cellular nucleus, neatly walled offfrom the various or-
ganelles that dot the plains of their rypical cell's cytoplasm. It was
the "not all" that attracted her attention: some of these extra-nuclear
organelles-specifically, the power plants of all animal and plant cells,
 Niles Eldredge

the "mitochondria"-were also known to have their own DNA.

ln plants, both mitochondria and chloroplasts, the locus of photo-
synthesis, have their own DNA complements. The sineple quesrion
she faced was: why? Why is there an independent set of genes in
these cytoplasmic organelles, when all of the "normal" genetic ma-
terial is otherwise organized as double sets of chromosomes within
the bounds of the nuclear walls?
Biological structures are signals of ancient evolutionary events.
'We owe the five fingers on our hands not to novel evolutionary
events a million years ago on the African savannas, but rather to
the original complement of five digits on the forefoot of the ear-
liest land vertebrates ("tetrapods"), who evolved some 370 million
years ago.
So, too, is mitochondrial DNA a holdover, a signal of an evolu-
tionary event. But this was like no other event ever proposed in evo-
lutionary annals: Lynn Margulis, to her everlasting credit, saw that
separate DNA complements imply the fusion of at least nvo differ-
ent kinds of other organisms, each with its own DNA complement,
to form a single, complex "eukaryotic" cell. Initially condemned as
heresy, this elegant idea had so much going for it that the biologi-
cal world has long since accepted it. There is simply no other plau-
sible explanation for the existence of separate DNA complements
in a "single" cell.
ln What Is Lfe?Lynn Margulis and Dorion Sagan tell us precisely
which kinds of bacteria fused to form the original nucleated cells-
olr cells. But that is far from all, for the Margulis mind, ever restless,
has kept on pushing the envelope. What Is Life? presents the case for
an even earlier evolutionary fusion of bacteria species. Margulis has
come to be convinced that such symbiotic origins of novel life forms
("symbiogenesis") has been far more cofiunon than ever dreamt by
evolutionary biologists steeped in the Darwinian tradition-a tra-
dition that emphasizes competition far more than cooperation in
the evolutionary process. Symbiogenesis is Margulis's central con-
tribution to the evolutionary dialogue, which has become enriched
 xtv Foreword

through her efforts to see the grand implications latent in the his-
tory of the microbial world.
But there is more to the Margulis-Sagan canon than even these
profoundly new, and heretofore undreamt, philosophies. Tireless
champions of the microbial world, the authors have labored might-
ily in an almost public-relations sense, striving to reveal the im-
mensely diverse array of microorganisms. For rnicrobes will not only
inherit the earth (should, for example, we complex multicellular
creatures fall prey to the next spasm of mass extinction); microbes
got here long before we did, and in a very real sense they already
"own," and most certainly run, the global system. They fix and re-
cycle nitrogen and carbon and other essential elements otherwise
unavailable to our bodies; they produce oxygen, natural gas
(methane), and so on and on. Without the microbial world, liG as

we ourselves experience it simply could not be.

All of which lifts the Margulis gaze from the microscopic to the
global: Earth truly is a living system, a globally pulsing amalgam of
organisms and the physical "inanimate" world. Whether or not one
chooses to call this system "Gaia" and pronounce it as alive as any
organism does not, in a profound sense, really matter. For in read-
ing What k Life? we see, clearly and simply, that the global systenl
linking liG with the physical realm truly does exist, and that we hu-
mans, despite appearances and protestations to the contrary, are still
very much a part of that system.
Which takes us back to the ultimate value of being aware of our
own existence. As we read What Is L!fe?, we think about life's ri-
otous diversity and evolution's exuberance, and we realize that the
global system, all that life, and, in the end, our very own existence,
are very much under threat-from our very own selves. What Is
Life? cornbines the stranger-than-fiction realities of the living world
with the kind of intellectual force that can reveal new undreamr
philosophies. It yields the understanding we so desperately need if
we are to confront the mounting threat we humans pose to the global
 Niles Eldredge ] xv

ecosystem as we cross over the millennial divide. Knowledge is

power, and What k LiJe? equips us with an understanding of the
living world that we so desperately need if we-along with the
world's ecosystems-are to survive.

Niles Eldredge
American Museum of Natural History
 LIFE: THE ETERNAL ENIGMA

Life is something edible, lovable, or lethal.

-,AMES E. LOVELOCK

Life is not a thing or a fluid any more than heat is. What we
observe are some unusual sets of objects separated from
the rest of the world by certain peculiar properties such as
growth, reproduction. and special ways of handllng energy.
These objects we elect to call "living things."
ROBERT MORISON

IN THE SPIRIT OF SCHRODINGER

Half a century ago, before the discovery of DNA, the Austrian

physicist and philosopher Erwin Schrodinger inspired a generation
of scientists by rephrasing for them the timeless philosophical ques-
tton: What ls Life? (fig. ,). In his classic 1944 book bearing that ti-
tle, Schrodinger argued that, despite our "obvious inabfity" to define
it, life would eventually be accounted for by physics and chemistry.
Life, Schrodinger held, is matter which, like a crystal-a strange,
"aperiodic crystal"-rspeats its structure as it grows. But life is far
more fascinating and unpredictable than any crystallizing mineral:

The difference in structure is of the same kind as that berween an or-

dinary wallpaper in which the same pattern is repeated again and agun
in regular periodiciry and a masterpiece of embroidery, say a Raphael
tapestry, which shows no dull repetition, but an elaborate, coherent,
meaningful design traced by the great master.l
 What ls Life?

FICURE 1. Erwin Schrodinger: a physicist

whose emphasis on the physiochemical
nature of life helped inspire the discov-
ery of DNA and the molecular biological
revolution.

Schrcidinger, a Nobel laureate, revered life in all its marvelous com-

plexiry. Indeed, although he devised the wave equarion that helped
give quantum mechanics theory a firm mathematical basis, he never
conceived of life as simply a mechanical phenomenon.
Our book, addressing life's fullness without sacrificing any sci-
ence, reproduces not only Schrodinger's title but also, we hope, his
spirit. We have tried to put the life back into biology.

What is life? is surely one of the oldesr quesrions. We live. We-

people, birds, flowering plants, even algae glowing in the ocean at
night-differ from steel, rocks, inanimate matter.
-We
are alive. But what does it mean to live, to be alive, to be a
discrete being at once part of the universe but separated from it by
our skin? What is life?
Thomas Mann (r 8 7 5- r 9 5 5) gave an admirable, if literary, answer
in the novel The Magk Mountain:
'What
was life? No one knew. It was undoubtedly aware of itself, so
soon as it was liG; but it did nor know what it was . . . it was not mar-
ter and it was not spirit, but something berween the rwo, a phenom-
enon conveyed by matter, like the rainbow on the waterfall, and like
the flame. Yet why not marerial?-ir was senrienr to the point of de-
 Life: The Eternal Enigma I a

sire and disgust, the shamelessness of matter become sensible of it-

self, the incontinent form of being. It was a secret and ardent stirring
in the frozen chastity of the universal; it was a stolen and voluptuous
impuriry of sucking and secreting; an exhalation of carbonic gas and
material impurities of mysterious origin and composition.2

Our ancestors found spirits and gods everywhere, animaring all of

nature. Not only were the trees alive but so was the wind howling
across the savanna. Plato, in his dialogue l-aws,'said that those per-
Gct beings, the planets, travel around Earth voluntarily in circles.
MedievalEuropeans believed the microcosm, the small world of the
person, mirrored the macrocosm, the universe; both were part mat-
ter and part spirit. This ancient view lingers in the animals of the
zodiac and in the astrological notion that celestial bodies influence
mundane ones.
In the seventeenth century the German astrologer-astronomerJo-
hannes Kepler (r 57r-r63o) calculated that planets including Earth
travel around the sun in ellipses. Nevertheless, Kepler (who wrote
the first work of science fiction and whose mother was arrested as
a witch) believed that the stars inhabit a three-kilometer-thick shell
far beyond the solar system. He considered Earth a breathing, re-
membering, habit-forming monster. Although Kepler's view of a
living Earth now seems whimsical, he reminds us that science is as-
ymptotic: it never arrives at but only approaches the tantalizing goal
of final knowledge. Astrology gives way to astronomy; alchemy
evolves into chemistry. The science of one age becomes the mythol-
ogy of the next. How will future thinkers assess our own ideas? This
movement of thought-of living beings questioning themselves and
their surroundings-is at the heart of the ancient question of what
it means to be alive.

Life-from bacterium to biosphere-maintains by making more of

itself. We focus on self-maintenance in our first chaPter. Next, in
chapter 2, we trace views of life from very early on through Euro-
 4 What ls Life?

pean mind-body dualism and then to nlodern scientific material-

ism. Chapter 3 explores life's origins and its memory-like preser-
vation of the past. Our ancestors-the bacteria that brought Earth's
surface to life-are Gatured in chapter 4.
Through symbiotic mergers, bacteria evolved into the protists of
chapter 5. Protists are unicells, including algae, amebas, ciliates, and
other postbacterial cells with eroric habits anticipating our own:
they evolved into multicelled beings experiencing sex and death.
We call the unicellular protists, together with their close multicel-
lular relatives-some of which are very large-protoctists. The bac-
teria that formed protoctists were to have a spectacular future. They
became animals (chapter 6), fungi (chapter 7), and plants (chaprer
8). In the last chapter we pursue the unorthodox but commonsen-
sical idea that life-not just human life but all life-is free to act
and has played an unexpectedly large part in its own evolution.

LIFE'S BODY

Life, although material, is inextricable from the behavior of the liv-

ing. Defying definition-a word that means "to fix or mark the lim-
its of"-living cells move and expand incessantly. They overgrow
their boundaries; one becomes rwo become many. Although ex-
changing a great variery of materials and communicating a huge
quantity of information, all living beings ultimately share a com-
mon past.
Perhaps even more than Schrodinger's "aperiodic crystal," life re-
sembles a fractal-a design repeated at larger or smaller scales. Frac-
tals, beautiful for their delicacy and surprising in their apparent com-
plexiry are produced by computers, as graphics programs irerare, or
repeat, a single mathematical operation thousands of times. The
"fractals" of life are cells, arrangements of cells, many-celled or-
ganisms, communities of organisms, and ecosystems of communi-
ties. Repeated millions of times over thousands of mrllions of years,
the processes of life have led to the wonderful, three-dimensional
patterns seen in organisms, hives, cities, and planetary life as a whole.
 Life: The Eternal Enigma

Life's body is a veneer of growing and self-interacting matter en-

casingEarth. Twenty kilometers thick, its top is the atmosphere and
its bottom is continental rock and ocean depths. Life's body is like
a tree trunk. Only its outermost tissues grow. lJnless protected by
technology, itself an extension of life, any individual removed from
the living sphere is doomed.
Life, as far as is known, is limited to the surface of this third planet
from the sun. Moreover, living matter utterly depends on this sun,
a medium-sized star in the outback of the Mrlky'Way Galaxy. Less
than one percent of the solar energy that strikes Earth is diverted
to living processes. But what life does with that one percent is as-
tounding. Fabricating genes and offspring from water, solar energy,
and air, festive yet dangerous forms mingle and diverge, transform
and pollute, slaughter and nurture, threaten and overcome. Mean-
while, the biosphere itself, subtly changing with the comings and
goings of individual species, lives on as it has for more than 3,ooo
million years.

ANIMISM VS. MECHANISM

If you wish to, you can reach for a glass of water or snap this book
shut. From the experience of willing our bodies to move came
animism: the view that winds come and go, rivers flow, and ce-
lestial bodies guard the heavens because something inside each wills
the movement. In animism all things, not only animals, are seen
to be inhabited by an inner, animating spirit. Formalized in poly-
theistic religion, the multipliciry of gods-a moon god, Earth god,
sun god, wind god, and so on-was replaced in Islam, Judaism,
and Christianity by a single god who crafted the world. W.inds and
rivers and celestial bodies lost their will, but living organisms-
especially humans-retained theirs.
Finally, the last outposts of animism-living organisms-yielded
to the philosophy of mechanism. Motion need not imply any in-
ner consciousness; the program could have been "built in" by a cre-
ator. Wind-up toys and automated models of the solar system sug-
 What ls Life?

gested to their inventors that even living things may be constructible

from liGless mechanisms, subtle concealed springs, tiny unseen pul-
leys, levers, cogs, and gears. Comparing flowing blood to a hydraulic
system, the heart to a pump, English physician 'William Harvey
Q 578*1647) discovered circulation of the blood. Scientists sleuthed
out the world's secret mechanisms, part of an overall design. Nat-
ural history revealed the world to be a giant mechanism made ac-
cording to the mind of an omnipresent, omnipotent god.
Isaac Newton (t642-r727) became the high priest of mechanism.
A devoted student of alchemy, scripture, and the occult, Newton
made unparalleled innovations in optics, physics, and mathematics.
In doing so he helped bridge the gap from the medieval cosmos to
the modern one. Explaining the motions of the planets with a new
law of graviry Newton's equations showed that the world of the
heavens and that of Earth were one and the same; the force that
kept the moon in orbit was also the force that thuds an apple to the
ground. So revealing were Newton's discoveries of "laws" govern-
ing the entire universe that to some it seemed he had-in Kepler's
words-"glimpsed the mind of God." Inspired by Newton's analy-
ses, Pierre-Simon de Laplace Gl4g-r827) speculated that, with
sufilcient information, the entire future of the universe, even the
most minute human action, could be predicted. Far from being
moved by hidden spirits, the celestial bodies now seemed to be un-
der the governance of preexistent mathematical laws. Divine in-
tervention became increasingly superfluous. God did not need to
fiddle with creation. He had crafted it to last. The cosmos worked
itself.
'With a grasp of gravitation's
cosmic sweep, scientists were spurred
to explore phenomena once considered beyond human compre-
hension. Electriciry and magnetism, sound and colors, radiation and
heat, explosion and chemical change were all described with an eye
to their underlying unity. Optical instruments, telescope and mi-
croscope, presented formerly unseen worlds of the very far and the
very near. Experiment and criticism replaced blind acceptance of
 Life: The Eternal Enigma I Z

classical authoriry and divinely revealed truth. Scientists coaxed na-

ture to yield some of her most private secrets. Oxygen's role in fire,
lightning as electrical discharge, graviry as the invisible force caus-
ing the tides and attracting the moon into Earth's orbit-one by
one nature laid down her cards.
lJnder the spell of the mechanical worldview, the ancient al-
chemical dream of shaping nature to human will became techno-
logical realiry. After centuries of humans meddling with steamy con-
coctions in a Faustian quest to be godlike, then a 1953 discovery
seemed to reveal the very secret of life. Life was chemical and the
material basis of herediry was DNA, whose helical and staircase-
like structure made clear how molecules copied themselves. Indeed,
the "aperiodic crystal" that Schrodinger had predicted was uncan-
nily similar to the double helix first described by theEnglish chemist
Francis Crick and American whiz kid James D. 'Watson. Replica-
tion was no longer beholden to a mysterious "vital principle"; it
was the straightforward result of interacting molecules. The de-
scription of how DNA fabricated a copy of itself out of ordinary
carbon, nitrogen, and phosphorus atoms was perhaps the most spec-
tacular of all mechanism's successes. But paradoxically, this success
born of self-directed minds seemed to portray life-including the
scientists themselves-as the result of atoms involuntarily interact-
ing according to changeless and inviolable chemical law.
Bet'oveen these rwo extremes-the entire universe as alive, and
the living organism as chemical and physical machine-lies the
panorama of opinion. But is there not something wrong with both
the mechanizatron of life and the vitalization of matter?
The world as a vast machine fails to account for our own self-
awareness and self-determination because the mechanical world-
view denies choice. Mechanisms, after all, don't act; they react. And
mechanisms, moreover, don't come into existence on their own.
The assumption that the universe is a mechanism implies that it
was made according to some humanlike design-that is, by some
living creator. In other words, successful as it is, the scientific mech-
 8 I what ts Life?

anistic worldview is deeply metaphysical; it is rooted in religious

assumptions.
The animistic view of the cosmos as a huge organism is also
flawed. It blurs the distinctions among what is living, what is dead,
and what has never been alive. If everything were alive, there would
be no interest in-and scientists never would have discovered the
replicative chemistry of-lifb.
We thus reject mechanism as naive and animism as unscientific.
Even so, life, as an emergent behavior of matter and energy, is best
known by science. Schrcidinger was correct in advocating a search
for the physicochemical underpinnings of life. So are'Warson and
Crick and other physicists and molecular biologists who hail the
structure of DNA as a key to liG's secrets. Like an uncoiling spring
pushing the soft gears of life, DNA copies itself as it directs the mak-
ing of proteins that together form the leopard's spots, the spruce
tree's cone, and living bodies in general. lJnderstanding how DNA
works may be the greatest scientific breakthrough in history. None-
theless, neither DNA nor any other kind of molecule can, by itself.
explain life.

JANUS AMONG THE CENTAURS

The American architect R. Buckminster Fuller (r 895-r983) applied

"synergy" (from Greek synergos, working together) to describe en-
tities that behave more than the sum of their parts. From a sci-
as

entific standpoint, life, love, and behavior appear to be synergistic

phenomena. When certain chernicals-in water and in oil-came
together long ago, life was the result. Synergy also fits the emer-
gence of protist cells from bacteria, and of animals from such cells.
The common view is that life evolves by random genetic change
that is, moreover, detrimental more often than not. Chance muta-
tions, blind and undirected, are touted as the leading source of evo-
lutionary novelry. We (and a growing contingent of like-minded
students of life) do not entirely agree. Great gaps in evolution have
 Life: The Eternal Enigma ] S

been leaped by symbiotic incorporation of previously refined

components-components that have been honed in separate line-
ages. Evolution doesn't start anew each time a new life form appears.
Preexisting modules, which turn out to be primarily bacteria, al-
ready generated by mutation and retained by natural selection, come
together and interface. They form alliances, mergers, new organisms,
whole new complexes that act and are acted on by natural selection.
But natural selection by itself cannot generate any evolutionary
innovation, as Charles Darwin (r 8o9-r 892) was well aware. Natural
selection, rather, relentlessly preserves the former refinements and
newly generated novelry by culling those less able to live or re-
produce. Biotic potential-life's tendency to reproduce as much as
possible-takes care of the rest. But first, novelty must arise from
somewhere. [n synergy nvo distinct forms come together to make
a surprising new third one.
Cowboys, for example, settled the American-West. Some na-
tive Americans perceived the human-horse invaders as centaurs-
two-headed, multilimbed beings. The novelist and philosopher
Arthur Koestler (r9o5-r983) has called the coexistence of smaller
beings in larger wholes "holarchy.": Most people, by contrast, think
that life on Earth is hierarchical, a great chain of being with humans
on top. Koestler's coinage is free of implications of "higher" or of
one of the constituents in the holarchy somehow controlling the oth-
ers. The constituents, too, were given a new name by Koestler. Not
merely parts, they are "ho1ons"-wholes that also function as parts.
In his metaphysical as well as terminological rethinking, Koestler
invoked the double-facedJanus, who in Roman mythology was the
god of portals and the patron of beginnings and endings. In our
view, just as Janus simultaneously looks backward and forward, so
humans are not at the height of creation but point dually to the
smaller realnr of cells and the larger domain of biosphere. Life on
Earth is not a created hierarchy but an emergent holarchy arisen
from the self-induced synergy of combination, interfacing, and re-
cornbination.
 What ls Life?

BLUE JEWEL

The best part of a journey can be returning. By sending monkeys

and cats into orbit, people to the moon, and robots to Venus and
Mars, humankind has developed a new respect for, and a deeper
understanding of, life on Earth.
In r96r the Soviet lJnion's Vostok I carried the first human into
orbit around Earth. Since then, gazing "down" at this turquoise
orb-venturing out on a spacewalk as if about to jump from the
world's highest diving board-cosmonauts and astronauts have
groped for words that do justice to their experience. Eugene A. Cer-
nan, an astronaut of both the Gemini and Apollo lunar missions,
and the last person to walk on the moon, describes the view:

When you are inEarth orbit lookrng down you see lakes, rivers, penin-
sulas.. . . You quickly fly over changes in topography, like the snow-
covered mountains or deserts or tropical belts-all very visible. You
pass through a sunrise and sunset every ninefy rninutes. 'When you
leaveEarth orbit . . . you can see frompole to pole and ocean to ocean
without even turningyour head. . . . You literaily see North and South
America go around the corner Earth turns on an axis you can't see
as

and then miraculously Australia, then Asia, then all of Arnerica comes
to replace them. . . . You begin to see how little we understand of
time. . . . You ask yourself, where am I in space and time? You watch
the sun set over America and rise again over Australia. You look back
"home" . . . and don't see the barriers of color, religion, and poli-
tics that divide up this world.a

Imagine yourself in orbit. As you circle the planet every ninety min-
utes, time and space undergoes a mutual metamorphosis. Graviry
lessens;north and south become relafive. Day follows night in a patch-
work blend. The sun cuts through the thin ribbon that is the at-
mosphere, flooding the cabin of the spacecraft with red to green to
purple, through all the colors of the rainbow. You are plunged into
black. Earth becomes the place where there are no stars. If Earth can
be seen at all it is as a flicker of tiny lights-cities-on the surface
 Lif e: The Eternal Enigma | ,l.|

of the sun-eclipsing globe. "Day" breaks again, revealing the cloud-

flecked blue ocean. As you are jettisoned into hyperperspective, the
sky is now below. As if floating dreamily away from your own body,
you watch the planet to which you are now tied by oniy the invisi-
ble umbilical cord of graviry and telecomrnunication.
The act of viewingEarth from space echoes that of a baby glimps-
ing, and really seeing, itself in a mirror for the first time. The as-
tronaut gazes upon the body of life as a whole. The French psy-
choanalystJacques Lacan posits a stage in human development called
"the rnirror stage."s The infant, unable to control its limbs, looks
into the mirror and perceives its whole body. Humaniry's jubilant
perception of the global environment represents the mirror stage
of our entire species. For the first time we have caught a glimpse of
our full, planetary form. We are coming to realize that we are part
of a global holarchy that transcends our individual skins and even
humaniry as a whole.
Television images h ry69 revealed astronauts bounding over the
lunar dust. The moon, once a synonym for the unattainable, was
reached. A cratered wasteland, bone-dry, the moon was neverthe-
less still daunting in its lifelessness. As the cosrnic perspective was
broadcast, we homebodies were given a futuristic ride and were
offered a new view of the world, a new worldview with the power
to rally Earth's peoples around an icon more potent than any flag.
Members of disparate religious and spiritual traditions could now
join together as citizens of Earth. Individuals so affected, those who
saw the potential, came to know that the whole former under-
standing of life was parochial, a result of where we lived. Even time
was upset: night became shadow.
Tiibal conflicts, national politics, and the colored geographic re-
gions of maps are invisible from space. Science has, of course, re-
vealed to us that this blue jewel orbits but a lackluster star in the
outskirts of a spiral galaxy with myriad stars within a universe of
myriad galaxies. Al1 our history and civilization has transpired un-
der the gaseous blanket of, really, a middling planet in one solar sys-
 12 What ls Life?

tem. Voyaging in space, we saw Earth as home. But it is more than

home: it is part of us. In contrast with the pale moon in the dead
solar system of our galactic suburbs, this third planet from the sun,
our Earth, is a blue-and-white flecked orb that looks alive.

IS THERE LIFE ON MARS?

unexpectedly, the search for life on Mars provided scientific con-

firmation of the "body" of life as a whole on Earth. The Viking
mission, launched in ry75, sent tvvo orbiters and rwo landers to
Mars. Although returning spectacular images of "Marscapes," the
Viking landers performed a series of experiments that failed to find
any evidence of Martian 1ife. Channels carved by ancient rivers were
seen, fueling hopes that evidence for past life may yet be found on
the red planet.
One scientist, however, was able to search for life on Mars before
the Viking mission was launched. In 1967 James E. Lovelock, En-
glish inventor of a device that measures chlorofluorocarbons impli-
cated in the production of ozone holes, was consulted by the Na-
tional Aeronautics and Space Administration (NASA) in its search
for extraterrestrial life. NASA was interested in what Lovelock's in-
vention, a gas-measuring instrument some thousand times more sen-
sitive to certain atmospheric constituents than any previous device,
might reveal about Mars. An atmospheric chemist, Lovelock sus-
pected that, in principle, life on any planet could be detected by the
chemical markers left in that planet's air. Because the constituents
of Mars's atmosphere were already known by the spectroscopic sig-
nature of the planet's reflected light, Lovelock believed the data al-
ready sufiicient to determine whether Mars was a living planet. His
conclusion: Mars was devoid of life. Indeed, he boasted with his
own brand of quiet iconoclastic mischief that his prediction pre-
cluded any need to visit Mars at all and that he could save NASA a
prodigious sum of money.
Lovelock had measured Earth's atmospheric gases with a chro-
 Life: The Eternal Enigma | ,S

matograph outfitted with his new supersensitive "electron capture

device." He was startled: the chemistry of Earth's atmosphere, not
at all like the atmospheres of Mars and Venus, was utterly improb-
able. He found that methane, the chief constituent of natural gas
and present in the atmospheres of the four giant planets (Jupiter,
Saturn, (Jranus, and Neptune), freely coexisted in Earth's atmos-
phere with oxygen at concentrations more than ro3s times higher
than expected.
Methane exists at only one to tvvo parts per million in Earth's
atmosphere, but even that minuscule proportion is far too high.
Methane (one carbon atom surrounded by four hydrogen atoms)
and oxygen gas (two oxygen atoms) react explosively with each other
to generate heat, producing carbon dioxide and water. Oxygen, the
second most abundant gas in the atmosphere, should thus react im-
mediately with methane to make the latter undetectable. Perhaps in
the next minute you will die of asphyxiation because all the oxy-
gen atoms will gather in one corner of the room and your brain
will be deprived of its absolute requirement for oxygen gas. Such a
calarnity is improbable to the point of absurdiry. Yet the chemical
mixture of methane and oxygen in the Earth's air is equally freak-
ish. Indeed, not only methane but many other gases in our air
should not be detectable, given standard rules of chemical mixing.
Given their tendency to react with oxygen, some of our atmosphere's
components-methane, ammonia, sulfur gases, methyl chloride, and
methyl iodide-are far from chemical equilibrium. Carbon monox-
ide, nitrogen, and nitrous oxide are respectively ten, ten thousand
million, and ten trillion times more abundant than chemistry alone
can account for.
Biology, however, offers an answer. Lovelock realized, for instance,
that methane-producing bacteria release this gas in globally signi-
ficant amounts. Cows contribute methane by belching. Belched
methane does react with oxygen but, before it disappears, more is
produced. The methane is made from grass by bacteria and protists
in the cow's rumen, a special stomach.
 14 What ls Life?

Life has made our atmosphere chemically reactive and orderly,

while exporting heat and disorder to space. Lovelock maintained
that the atmosphere is as highly ordered as a painted tortoise's shell
or a sand castle on a deserted beach. And life's inveterate ordering
has left its traces on other planets. On zo July ry76 a lander was left
on Mars by the 3.6-metric-ton Viking I spacecraft. Although not
what scientists were looking for, this machine, sitting 57r rnillion
kilometers away at Chryse Planitia on red sand, is the best, and so
far the only evidence of life on Mars: solar-system exploring, tech-
nological human life.

LIFE A5 VERB

Lovelock's analyses have pushed biologists to realize that life is not

confined to the things now called organisms. Self-transforming, ho-
larchic life "breaks out" into new forms that incorporate formerly
self-sufiicient individuals as integral parts of greater identities. The
largest of these levels is the planetary layer, the biosphere itself. Each
level reveals a different kind of "organic being." This is the term
that Darwin used throughout his opus, Orz the Origin of Species. (" Or
ganism," like "scientist" and "biology," had not yet been coined.)
"Organic being" merits resurrection as it affords the recognition that
a "cell" and the "biosphere" are no less alive than an "organism."
Life-both locally, as animal, plant, and microbe bodies, and glob-
ally, as the biosphere-is a most intricate material phenomenon. Life
shows the usual chemical and physical properties of matter, but with
a fwist. Beach sand is usually silicon dioxide. So are the innards of
a mainframe computer-but a computer isn't a pile of sand. Life is
distinguished not by its chemical constituents but by the behavior
of its chemrcals. The question "'What is life?" is thus a linguistic trap.
To answer according to the rules of grammar, we must supply a
noun, a thing. But life on Earth is more like a verb. It repairs, main-
tains, re-creates, and outdoes itself.
This surge of activiry which not only applies to cells and animals
 Life: The Eternal Enigma 15

but to Earth's entire atmosphere, is intimately connected to two of

science's most famous laws-the laws of thermodynamics. The first
law says that throughout any transformation the total energy of any
system and its environment is neither lost nor gained. Energy-
whether as light, movement, radiation, heat, radioactiviry chemi-
cal or other-is conserved.
But not all forms of energy are equal; not all have the same efGct.
Heat is the kind of energy to which other forms tend to convert, and
heat tends to disorganize matter. The second law of thermodynam-
ics says that physical systems tend to lose heat to their surroundings.
The second law was conceived during the Industrial Revolution,
when the steam engine represented the state-oGthe-art in engi-
neering. French physicist Nicolas Carnot Q796-r832), aiming to
improve the efiiciency of the steam engine (whose governor mech-
anism was invented byJames Watt [r736-1 8 r9]), came to realize that
heat was associated with the movement of minute particles. And
from that, he envisioned the principle that is now known as the sec-
ond law: In any moving or energy-using system entropy increases.
In systems undergoing change, such as steam engines or electric
motors, a certain amount of the total energy available is already in,
and more is converted into, a form that is unavailable for useful work.
Although the amount of energy in the system and its environment
stays the same (i.e., the first law of thermodynamics, of conserva-
tion of energy, holds), the amount of energy available to do work
decreases. In computer science entropy is measured as the uncer-
tainry in the information content of a message. The second law un-
equivocally claims that in changing systems entropy increases. im-
plying that heat, noise, uncertainry and other such forms of energy
not useful for work, increase. As local systems lose heat, the uni-
verse as a whole is gaining it. Although not so popular now, in the
past physicists and chemists have made the prediction that the uni-
verse will whimper out in a "heat death" as a consequence of the
tendency for entropy to increase. More recently, they have even in-
vented the',vord "negentropy" for life, which, in its tendency to in-
 16 What ls Life?

crease information and certainry seems to contradict the second law.

It doesn't; the second law holds as long as one regards the system
(life) in its environneent.
In steam engines, coal was burned and carbon joined with oxy-
gen, reaction that, generating heat, made machine parts move. The
a

leftover heat that was generated was unusable. The heat in a cabin
on a snow-covered mountain seeks with seenring purpose any avail-
able crack or opening to mix with the cold air outside. Heat natu-
rally dissipates. This dissipative behavior of heat illustrates the sec-
ond law: the universe tends toward an increase in entropy, toward
even temperatures everyrvhere, as all the energy transforms into use-
less heat spread so evenly that it can do no work. Heat dissipation,
we are usually told, results from random particle motion. But there
are other interpretations.
Some scientists have begun to interpret the second law's predilec-
tion for heat-energy as the basis for apparent purposeful action. Ilya
Prigogine, a Belgian Nobel laureate, helped pioneer the consider-
ation of life within a larger class of "dissipative structures," which
also includes decidedly nonliving centers of activity like whirlpools,
A rather awkward term because it focuses
tornadoes, and flames.6
on what the structures-actually, systems, not structures-throw
away rather than what they retain and build, a dissipative system
maintains itself, and nlay even grow, by importing "useful" forms of
energy and exporting, or dissipating, less useful forms-notably, heat.
This thermodynamic view of life actually goes back to Schrcidinger,
who also likened living beings to flames, "streams of order" that
maintain their forms.
American scientist Rod Swenson has argued that the seeming pur-
with time is intimately
pose displayed in heat's tendency to dissipate
related to the behavior of life forms striving to perpetuate them-
selves.In Swenson's view, this entropic universe is pocked by local
regions of intense ordering, including life, because it is through or-
dered, dissipative systems that the rate of entropy production in the
universe is maximized. The more life in the universe, the faster that
various forms of energy are degraded into heat.7
 Life: The Eternal Enigma I l,

Swenson's view shows how life's seeming purpose-its seeking

behavior, its directedness, which philosophers cal, teleology-is re-
Iated to the behavior of heat. Scientists do not as a rule endorse
teleology. They consider it unscientific, a holdover from the prim-
itive days of anirnism. Teleology is nevertheless embedded in lan-
guage, and it cannot and need not be eliminated from the sciences.
The prepositions "to" and "for," which build teleology-that is,
purposefulness-into language, speak of a future-directedness that
seems present, to some degree, in all living beings. One should not
assume that only huneans are future-oriented. Our own frenetic at-
tempts, and those of the rest of life, to survive and prosper are a
special, 4,ooo-nrillion-year-old way the universe has organized it-
self "to" obey the second law of thermodynamics.

SELF-MAINTENANCE

Islands of order in an ocean of chaos, organisms are far superior to

human-built machines. Unlike James'Watts's steam engine, for ex-
ample, the body concentrates order. It continuously self-repairs.
Every five days you get a new stomach lining. You get a new liver
every r'rvo months. Your skin replaces itself every six weeks. Every
year, 98 percent of the atoms of your body are replaced. This non-
stop chendcal replacement, metabolism, is a sure sign of life. This
"machine" demands continual input of chemical energy and ma-
terials (food).
Chilean biologists Humberto Maturana and Francisco Varela see
in metabolism the essence of something quite fundamental to life.
They call it "autopoiesis." Coming from Greek roots meaning self
(auto) andmaking (poiein, as in "poetry"), autopoiesis refers to life's
continuous production of itself.8 Without autopoietic behavior, or-
ganic beings do not self-maintain-they are not alive.
An autopoietic entiry metabolizes continuously; it perpetuates
itself through chemical activiry the movement of molecules. Au-
topoiesis entails energy expenditure and the making of messes. Au-
topoiesis, indeed, is detectable by that incessant life chemistry and
 18 What ls Life?

energy flow which is metabolism. Only cells, organisms made of

cells, and biospheres made of organisms are autopoietic and can
metabolize.
DNA is an unquestionably importanr molecule for life on Earth,
but the molecule itself is not alive. DNA molecules replicate but
they don't metabolize and they are not autopoietic. Replication is
not nearly as fundamental a characteristic of life as is autopoiesis.
Consider: the mule, offspring of a donkey and a horse, cannot "repli-
cate." It is sterile, but it metabolizes with as much vigor as either of
its parents; autopoietic, it is alive. Closer to home, humans who no
longer, never can, or simply choose not to reproduce can not be
relegated, by the strained tidiness of biological definition, to the
realm of the nonliving. Of course, they too are alive.
In our view viruses are not. They are not autopoietic. Too small
to self-maintain, they do not metabolize. Viruses do nothing until
they enter an autopoietic entiry: a bacterial cell, the cell of an ani-
mal, or of another live organism. Biological viruses reproduce within
their hosts in the same way that digital viruses reproduce within
computers. Without an autopoietic organic being, a biological virus
is a mere mixture of chemicals; without a computer, a digital virus
is a mere program.
Smaller than cells, viruses lack sullicient genes and proteins to
maintain themselves. The smallest cells, those of the tiniest bacteria
(about one ten-millionth of a meter in diameter) are the minimd
autopoietic units known today. Like language, naked DNA mole-
cules, or computer progralrls. viruses mutate and evolve: but, by
themselves, they are at best chemical zombies. The cell is the small-
est unit of life.
When a DNA molecule produces another DNA moiecule ex-
actly like itself, we speak of replication. When living matter, as a
cell or as a body made of cells, grows another similar being (with
differences attributable to mutarion, genetic recor-nbination, sym-
biotic acquisition, developmental variation, or other factors), we
speak of reproduction (see plate z). When living matter continues
 Life: The Eternal Enigma I ,g

to reproduce altered forms that, in turn, make altered offspring, we

speak of evolution: change in populations of life forms over time.
As Darwin and his legacy stress, more reproducing cells and bodies
are produced by budding, cell division, hatching, birth, spore for-
mation, and the like, than can ever survive. Those that cope long
enough to reproduce are "naturally selected." Put more bluntly, it
is not so much that survivors are selected for their success as that
those who fail to reproduce before dying are selected against.
Identity and self-maintenance require metabolism. Metabolic
chernistry (often called physiology) precedes reproduction and evo-
lution. For a population to evolve, its members must reproduce. Yet
before any organic being can reproduce, it must first self-maintain.
'Within
the lifetime of a cell, each of five thousand or so different
proteins will completely interchange with the surroundings thou-
sands of times. Bacterial cells produce DNA and RNA (nucleic
acids), enzyme proteins, fats, carbohydrates, and other complex car-
bon chemicals. Protoctist, fungi, animal, and plant bodies all pro-
duce these and other substances as well. But most importantly, and
amazingly, any living body produces itself.
This energetic maintenance of uniry while components are con-
tinuously or interrnittently rearranged, destroyed and rebuilt, broken
and repaired, is metabolism, and it requires energy. In accordance with
the second law of thermodynamics, autopoietic self-maintenance
preserves or increases internal order only by adding to the "disor-
der" of the external world, as wastes are excreted and heat is vented.
All living beings must metabolize and therefore all must create 1o-
cal disorder: useless heat, noise, and uncertainry. This is autopoietic
behavior, reflecting the autopoietic imperative required for any or-
ganic being that lives, that continues to function.
The autopoietic view of life differs from standard teachings in bi-
ology. Most writers of biology texts imply that an organism exiss apart
from its environment, and that the environment is mostly a staric, non-
living backdrop. Organic beings and environment, however, inter-
weave. Soil, for example, is not unalive. It is a mixture of broken rock,
 20 What ls Life?

pollen, fungal filaments, ciliate cysts, bacterial spores, nematodes, and

other microscopic animals and their parts. "Nature," Aristode ob-
served, "proceeds little by little from things Lifeless to animal life in
such a way that it is impossible to determine the exact line of de-
marcation."e Independence is a political, not a scientific, term.
Since life's origin, all living beings, directly or circuirously, have
been connected, as their bodies and populations have grown. In-
teractions occur. as organisms connect via water and air. Darwin. in
his Origin of Species,likened the complexiry of these interactions to
"an entangled bank"-too complex for us humans even to begin
to sort out: "Throw up a handful of feathers, and all fall to the
ground according to definite laws; but how simple is the problen-r
where each shall fall compared to that of the action and reaction of
the innumerable plants and animals." Yet it is the sum of these un-
countable interactions that yields the largest level of life: the blue
biosphere, in all the holarchic coherence and mysterious grandeur
of its evolution from the black cosmos.

THE AUTOPOIETIC PLANET

The biosphere as a whole is autopoietic in the sense that it main-

tains itself. One of its vital "organs," the atmosphere, is clearly
tended and nurtured. Earth's atmosphere, approximately one-fifth
oxygen, differs radically from that of Mars and Venus. The atmos-
pheres of these planetary neighbors are nine parts in ten carbon
dioxide; in Earth's atmosphere, carbon dioxide accounts for only
three parts in ten thousand. If Earth's biosphere were not made of
carbon dioxide-consuming beings (plants, algae, and photosynthetic
and methane-producing bacteria, among myriad other life forms),
our atmosphere would long ago have reached carbon dioxide-rich
chemical stabiliry and virtually every molecule capable of reacting
with another molecule would already have reacted. Instead, the com-
bined activities of autopoietic surface life have led to an atmosphere
in which oxygen has been maintained at levels of about 20 percent
for at least 7oo million years (fig. z).
 COz 96.5%

Nz 32%

Trace gases 0.2%

02 20.9%
Trace gases 0.07%

COz 0.O3%

N2 79Y"

co2 95%

Nz 2.7Y"

Trace gases 2.3%

FIGURE 2. Atmospheric comparison of Earth and its two planetary

neighbors. Note the comparatively high concentration of the ex-
plosive gas oxygen and the very low concentration of carbon diox-
ide on Earth. This atmospheric anomaly results from the incessant
activity of gas-exchanging organisms. The minute physiology of the
cell over geological time becomes magnified into the global physi-
ology of the biosphere.
 22 What ls Life?

Other evidence for life on a planetary scale comes from astron-

omy. According to standard astrophysical models of the evolution
of stars, the sun used to be cooler than it is now. The sun's lumi-
nosity has increased by 3 o percent or more since life began on Earth.
Living things can grow and reproduce only in a limited temperature
range within which water is liquid. Fossils of life more rhan 3,ooo
million years old confirm that ancient temperatures were not all that
dissimilar from those prevailing today; other geological evidence sug-
gests that liquid water was widespread on Earth at least 4,ooo rnil-
lion years ago. The increase in the luminosiry of the sun should have
dramatically increased the surface temperature of Earth since those
early times. Because no dramatic increase has occurred-indeed,
the trend may have been a cooling-it appears that the tempera-
ture of the entire biosphere has been self-maintained. By respond-
ing, life seems to have succeeded in cooling the planetary surface
to counter, or more than counter, the overheating sun. Mainly by
removing from the atmosphere greenhouse gases (such as methane
and carbon dioxide) which trap heat, but also by changing irs sur-
face color and form @y retaining water and growing slime), life re-
sponded, prolonging its own survival.
Oceanography provides still another glimpse of the body of life
as a whole. Chemical calculations suggest that salts should accumu-
late in the oceans to concentrations perilous to nonbacterial forms
of life. Salts, such as sodium chloride and magnesium sulfate, are
continuously eroded from the continents and carried into the
oceans by rivers. World oceans have, however, remained hospitable
to salt-sensitive organisms for at least 2,ooo rnillion years. Seafaring
microorganisms may therefore be sensing and stabilizing ocean acid-
iry and salinity levels on a global scale. How life removes salt frorn
marine waters is obscure. Perhaps salt concentrations too high for
most life are lowered, in part, by the vigorous pumping of sodium,
calcium, and chloride out of cells and, in part, by formation of evap-
orite flats. These encrusted fields are rich in sea salt and salt-loving
microbes. They often form behind lagoonal barriers made by ani-
 Life: The Eternal Enigma I ,,

mals such as corals or when shifting sands are trapped by the mucus
and slime formed by microbial communities. Continuous desalina-
tion, if it exists, may be part of a global physiology.
Some evolutionary biologists have suggested that Earth life in its
totaliry cannot constitute a living body, cannot be a living being,
because such a body could only have evolved in competition with
other bodies of the same sort-presumably, other biospheres' But,
in our view, autopoiesis of the planet is the aggregate, emergent
property of the many gas-trading, gene-exchanging, growing, and
evolving organisms in it. As human body regulation of temperature
and blood chemistry emerges &om relations among the body's com-
ponent cells, so planetary regulation evolved from eons of interac-
tions among Earth's living inhabitants.
Using the energy of sunlight, only green plants, algae, and cer-
tain green- and purple-colored bacteria can convert compounds
from surrounding water and air into the living stuffof their bodies.
This sun-energized process, photosynthesis, is the nutritional basis
for the rest of life. Animals, fungi, and most bacteria feed on the
purple and green producers. Photosynthesis evolved in microbes
soon after the origin of life. At every level, from microbe to planet,
organic beings use air and water or other organic beings to build
their reproducing selves. Local ecology becomes global ecology. As
a corollary, and in spite of English grarrunar, life does not exist o,?

Earth's surface so much as it rs Earth's surface.

Life extends over the planet as a contiguous, but mobile, cover and
takes the shape of the underlying Earth. Life, moreover, enlivens the
planet;Earth, in a very real sense, is alive. This is no vague philosophical
claim but rather a physiological truth of our lives. Organisms are less
self-enclosed, autonomous individuals than communities of bodies
exchanging matter, energy, and information with others. Each breath
connects us to the rest of the biosphere, which also "breathes," al-
beit at a slower pace. The biosphere's breath is marked daily by in-
creasing carbon dioxide concentrations on the dark side of the globe
and decreasing concentrarions on the lighted side. Annual breathing
3
'=
c
o
o-
E(d
o-

310 r
t"-- --*--.-+..*.-*-- --{-.. ----.------'*l
Years 1960 1970 1980 1990

FIcURE 3. Seasonal fluctuations of carbon dioxide in the northern hemisphere. The

peaks of the zigzags represent an increase in atmospheric carbon dioxide during sum-
mers; the overall upward trend indicates rising levels of COz due at least in part to
human activity. This seasonal and annual fluctuation of carbon dioxide in the Earth's
atmosphere attests to " breathing" on a global scale. The total carbon dioxide increase
may, by the greenhouse effect, raise planetary temperatures to levels inhospitable for
human beings-a geophysiological "fever."

is marked by the passage of the seasons; photosynthetic actrviry kicks

up in the northern hemispherejust as it is winding down in the south.

Taken at its greatest physiological extent, liG ls the planetary sur-

face. Earth is no more a planet-sized chunk of rock inhabited with
life than your body is a skeleton infested with cells (fig. :).

THE STUFF OF LIFE

When German chemist Friedrich Wohler (r8oo-i882) first, acci-

dentally, produced crystals of urea by heating ammonium cyanate,
he could not accept that he had made from scratch a compound so
clearly associated with living beings. lJrea, after all, is the carbon-
nitrogenous waste produced in animal urine. And in'W'cihler's day,
 Life: The Eternal Enigma I ,U

organic beings were believed to consist of a strange and wonderful

"organic matter" that was present in life-and nowhere else. Since
then, dozens of carbon-rich compounds, such as formic acid, eth-
ylene, and hydrogen cyanide, have been found notjust in life but
in interstellar The equivalent of an estimated ro quintillion
space.
(ro,ooo,ooo,ooo,ooo,ooo,ooo) fifths of whiskey, in the form of the
nine-atom molecule CH:CHzOH (ethyl alcohol), exists in one in-
terstellar cloud in the constellation of Orion alone.
Though adulterated with other compounds, we, like all living
matter, are mostly water-that is, hydrogen and oxygen. Hydrogen
forms, by mass, 7i percent of the atoms in the cosmos. It is the
same element which, under intense gravitational pressure, becomes
helium in the nuclear fusion reaction that makes our sun shine.
Far older and bigger stars went out with a bang, as supernovas, and
thereby created carbon, oxygen, nitrogen, and the other heavier
elements. Life is made from such star stuff. In the universe life
may be rare or even unique. But the stuff of which it is made is
commonplace.
More and more inert matter, over time, has literally come to life.
Minerals of the sea are now incorporated into living creatures for
protection or support in the form of integument, shell, bone. Our
own skeletons are built from calcium phosphate, a sea salt that was
initia[y a nuisance or a hazard for our remote ancestors' marine pro-
tist cells which eventually found ways to cleanse their tissues by put-
ting such minerals to use. The kinds as well as the mass of chemi-
cal elements in living bodies have increased through evolutionary
time. Whereas structural compounds made of hydrogen, oxygen'
sulfur, phosphorus, nitrogen, and carbon are required by all cells and
have been essential to life since its inception, those made of silicon
and calcium are relative newcomers.
Heinz Lowenstam (r9r3-r993), a Silesian-born geologist and
refugee from Nazi Germany, cataloged the minerals produced in the
hard parts of animals. In Lowenstam's youth, the only hard substances
thought to be produced by living tissues were the calcium phos-
phate of our own bones and teeth, the calcium carbonate of mollusk
FICURE 4. Oxalic acid crystal taken from a sea squirt renal sac, an organ
thought to be a ductless kidney. Nephromyces, a protoctist probably asso-
ciated with symbiotic bacteria, apparently forms the crystals from the ani-
mal's uric acid and calcium oxalate. Over fifty such minerals are now known
to be produced in living cells.
 Life: The Eternal Enigma I ,,

shells, and the silicon dioxide of unusual structures such as the spic-
ules of sponge. Lowenstam and his colleagues went on to discover
many other minerals produced by life, including calcium oxalate
crystals made by bacteria, plants, and others (fig. +).The list of hard
substances made in live cells, including unexpectedly beautiful crys-
tals, now surpasses fifry (table t).
Life had been reusing hard materials and shaping solid wastes long
before the appearance of technological humans. Bacteria came to-
gether to form protoctists that in turn could mine and use calcium,
silica, and iron from the world's seas. Protoctists evolved into ani-
mals with shells and bones. Animals, individually or in concert, en-
gineered inert materials into tunnels, nests, hives, dams, and the like.
Even some plants incorporate minerals. The silica-laced bodies of
"scouring rushes," for example, may serve as good pot scrubbers
for campers, but they have probably evolved to deter herbivores.
The calciune oxalate crystals of Diffinbachia are hurtled from the
leaf cells toward unwary, hungry victims.
The propensiry to "engineer" environments is ancient. Today
people make over the global environment. Clothed and bespecta-
cled inside an automobile, connected by phone wires and radio
waves to modems, cellular phones, and bank machines, supplied with
electriciry plumbing, and other utilities, we are transforming our-
selves from individuals into specialized parts of a global more-than-
human being. This metahuman being is inextricably bound to the
much older biosphere, from which it arose. Metals and plastics rep-
resent the newest realm of matter "coming to life."

MIND IN NATURE

The biological self incorporates not only food, water, and air-its
physical requirements-but facts, experiences, and sense impres-
sions, which may become memories. A1l living beings, not just an-
imals but plants and microorganisms, perceive. To survive, an or-
ganic being must perceive-it must seek, or at least recognize, food
and avoid environmental danger.
 MINERALS PRODUCED BY LIFE
KINCDOMS OF ORCANISMS
Minerals Bacteria Protoctista Fungi Animals plants

CALCIUM
Calcium carbonate sheath and ameba and extracellular corals; extracellular
(CaCO:; aragonite, other extracellular foraminiferan precipitates; mollusk shells; precipitates
calcite, vaterite) precipitates shells mushrooms echinoderm
skeletons;
calcareous
sponges;
some kidney
stones
Calcium phosphate extracellular brachiopod
(CaPOq) precipitates; " lamp shells" ;

mushrooms vertebrate
teeth and bones;
some kidney
stones
Calcium oxalate extracellular mostkidney Dieffenbachia,
(CaCzOa) precipitates stones a flowering
plant

stu coN
Silica (SiOz) precipitates diatom and glass-sponge grass
radiolarian spicules phytoliths;
shells; horse-tail
mastigote stems
algae scales

IRON

Magnetite(Fe3Oa) magnetosomes arthropods;

mollusks;
vertebrates
Greigite (Fe3S4) magnetosomes

Siderite (FeCO:) extracellular

precipitates

Vivianite extracellular
8H2O)
(Fe3[PO4]2 . precipitates

Goethite extracellular extracellular chiton

11f:-o_:9ll P:-:it't"t":
precipitates mollusks
Lepidocrocite extracellular extracellular chiton
(xxFeO.OH) precipitates precipitates; mollusks
mushrooms

Ferrihydrite mollusks flowering

(5FerO3 .9H2O) plants
 kontinued)
KINGDOMS OF ORGANISMS

Minerals Bacteria Protoctista Fungi Animals

MANCAN ESE

Manganesedioxide intracellularor
(MnOz) extracellular
precipitates
around spores

BARI UM

Barium sulfate algal-plastid sense orSans:

(BaSOr) gravity sensors; statoliths
marine protoc- (otoliths)
tist skeletons
(xenophyophores)

STRONTI UM

Strontium sulfate manne pro- mollusk shells

(SrSO+) tist shells
(actinopods)

TABLE '1 . Contrary to popular belief, minerals and am-

mds do not belong to separate kingdoms. Many minerals

are produced in and by life, sometimes in crysulline form.
One of the most common minerals, calcium carbonate, is
formed by living marine animals as shells. Another com-
pound, calcium phosphate, is precipitated by cells of our
bones. As this table shows, all five kingdoms of organisms
have members which produce minerals. This list represents
only a sample of the over fifty minerals now known to be
produced by living cells.
 30 What ls Life?

A living being need not be conscious ro perceive. But consider:

most of our own daily activities-breathing, digesting, even turn-
ing a page or driving a car-are performed largely or even wholly
unconsciously. From the viewpoint of the evolutionary biologist, it
is reasonable to assume that the sensitive, embodied actions of plants
and bacteria are partof the same continuum of perception and ac-
tion that culminates in our own most revered mental attributes.
"Mind" may be the result of interacting cells.
Mind is fully an evolutionary phenomenon. Hundreds of mil-
lions ofyears before organic beings verbalized life, they recognized
it. Discerning what could kill them, what they could eat, and whar
they could mate, roughly in that order, was crucial to animal sur-
vival. One lJ.S. Supreme Court justice avowed that while he might
not be able to define obsceniry he surely could recognize it when
he saw it. We all have a similar abiliry with life. Life has been rec-
ognizing itself long before any biology textbooks were wrimen.
Survival-based psychological tendencies infiltrate the pristine
realm of science. Pattern recognition was such a useful trait for our
ancestors that, even if occasionally wrong, the Aha!feeling of dis-
covery would have been reinforced. Aesthetic judgments of elegance
and beaury often cited in the preGrence for certain equations over
others in physics, show that scientific correctness can also be intu-
'What
itive. we know, what we are capable of knowing and seeing,
has been shaped by our evolution as surviving creatures. Even fool-
ish and outlandish notions would have been retained and reinforced
if they in any way aided our ancestors' survival.
Neuroscientists have traced subjective Gelings of pleasure to en-
dorphins and enkephalins, two groups of neuropeptides produced
by the brain. The pleasure associated with seeing beaury including
scientific "truth," may have come about during the course of evo-
lution, just as love and biophilia-the pleasure we rake in the com-
pany of other living creatures-provoke us to seek out mates and
the natural environments that have been most conducive to our sur-
vival.If we did nor fear death, we might be too quick to kill our-
selves when troubled or inconvenienced and thus perish as a species.
 Life: The Eternal Enigma I t't

Belief in life's importance may not be a reflection of realiry then,

but an evolutionarily reinforced fantasy that prejudices believers to
do what is necessary, bear whatever burdens, to survive.
We all inherit a shared perspective bequeathed by our ancestors.
The physicists' hope of solving an essential set of equations for all
time and the cosmos may be but the gleam of a receding mirage.
In the end, as Charles Peirce (r 839-r9r4) andWilliamJames (r 842-
rgro) recognized, there may be no better measure of "truth" than
that which works-that which helps us survive.
Mind and body, perceiving and living, are equally self-referring,
selirefledve processes already present in the earliest bacteria. Mind,
as well as body, stems from autopoiesis. And in sufiiciently expres-
sive humans the process of autopoiesis under\ing living organiza-
tion makes itself manifest even outside the body. Abstract expres-
sionist painter Willem de Kooning Q9o4-r997) wrote:

If you write down a sentence and you don't like it, but that's what
you wanted to say, you say it again in another way. Once you start
doing it and you find how dillicult it is, you get interested. You have
it, then you lose it again, and then you get it again. You have to change
to stay the same.l('

Changing to stay the same is the essence of autopoiesis. It applies

to the biosphere as well as the cell. Applied to species, it leads to
evolution.

so, wHAT lS LIFE? It is a material process, sifting and surfing over

matter like a strange, slow wave. It is a controlled, artistic chaos, a
set of chemical reactions so staggeringly complex that more than
eighry million years ago it produced the mammalian brain that now,
in human form, composes love letters and uses silicon computers
to calculate the temperature of matter at the origin of the universe.
Life, moreover, appears to be on the verge of perceiving for the first
time its strange but true place in an inexorably evolving cosmos.
Life, a local phenomenon of Earth's surface, can in fact be un-
 32 What ls Life?

derstood only in its cosmic milieu. It formed itself out of star stufl
shortly after Earth 4,6oo million years ago congealed from a rem-
nant of a supernova explosion. LiG may end in a mere roo million
years when, embattled by dwindling atmospheric resources and in-
creased heat from the sun, systems of global temperature regula-
tion finally fail.11 Or life, enclosed in ecological systems, may es-
' cape and watch from safe harbor as the sun, exhausting its hydrogen,
explodes into a red giant, boiling offEarth's oceans, 5,ooo million
years from now.
 LOST SOU LS

Ay, but to die, and go we know not where;

To lie in cold obstruction and to rot.
WILLIAM SHAKESPEARE

Love that endures for a breath:

Night, the shadow of light,
And life, the shadow of death.
ALGERNON SWINBURNE

DEATH: THE GREAT PERPLEXER

The scientific mystery of liG in a near-lifeless, mechanical universe

mirrors the enigma of death in a fully living, animistic one. Our
ancestors inhabited a world where warm, moving bodies would reg-
ularly stop, grow cold, and decay. As puzzling as life is for us, so was
death for them. But we moderns still feel the influence of ancient
solutions to the death puzzle.
Until the seventeenth century the sun and moon did not move
according to Newtonian principles; these celestial bodies often were
animated by spirits within them. The whistling of the wind, the
changing phases of the moon, the twinkling, turning stars-these
eternal, celestial bodies moved as they willed to move, as we move
by will. But what happened to the will of the warrior, whose heart
had beat so hotly a moment ago, and is now a cold corpse? Does
life slip away at spearpoint in a pool of blood? As the corpse be-
 34 What ls Life?

comes rigid, does the enlivening spirit dart into the grass? Vanish
into thin air?
Originally death, not life, was the great perplexer.

In a living universe what is death? Where do "we" go when we die?

A gold coin disappears in one hand as the magician produces an-
other just like it in the opposite. The spectator concludes the gold
migrated from hand to hand, just as the logical mind concludes that

the soul sneaks out of the body after death. Any nearby being could
have taken the missing soul. An infant, a goat, a snake-a raven at
the scene of the crime-could have snatched the essence whose lack
rendered a body lifeless.
An apparent attention to the mystery of death marks the earliest
human remains. Sixry thousand years ago, at Shanidar cave in lraq,
a Neanderthal man was buried on a mat of woven pine boughs and

with flowers related to grape hyacinth, bachelor's button, hollyhock,

and groundsel.l Such grave sites, filled with flower parts, pollen,
amulets, beads, headbands made of fox teeth, weapons, tools, and
food, attest to funerary rites seemingly designed to provide the soul
with rest-and with the goods it would require in an afterlife.

THE BREATH OF LIFE

No great leap of faith is needed to see how early mysteries of the

corpse led to religious notions of spirit. For the Iroquois of North
America, the spirit was an exceedingly refined image, possessing a
tiny body replete with head, teeth, and limbs. The Karo Battak of
Sumatra envisioned a "tendi," a copy of the owner or other self,
which flees at death. The peoples of Papua and Malay posited a
brown, corn kernel-sized semangat or semungi, which, if it departed
temporarily, induced malady and, if permanently, death. Even the
inventor of the microscope, Antoni vanLeeuwenhoek (tQz-t74),
thought he saw the homunculus-a tiny, human-shaped seed-
when he examined sperm.
 LostSouls I aS

Some cultures have regarded blood, others flesh (the Australians,

the kidney fat) as the seat of life. New Zealand Maoris held to the
notion that the menstrual blood is the source of life. Shadows, flames,
trees, columns, dolls, pools, children, and Polaroid photographs have
all been proposed to detain or permanently entrap souls.
Of the candidates for life's essence, breath is strongest. The an-
cient Chinese used durable, airtight cofiins of cypress and pine,
tightly binding and stuffing the mouths of their dead with jade, gold,
silver, pearls, and cowry shells to restrain the spirit. The word "spirit"
itself comes from spiritus, Latin for breath. Birth is announced by
crying-and breathing. So long as there is life, there is breath.
Breath is invisible. Like wind, it moves things. Moreover, we speak
with breath. Shamans and priests from many cultures concluded that
air as spirit, perhaps the holy spirit of some unseen breathing be-
ing, was the intangible link berween life and death. Witness the et-
ymological kinship between inspiration. expiration, respiration,
and spirit. Then too, expirarion is itself a synony'rn for death. In many
Native American languages Great Spirit and Great Wind share the
same word and meaning. The Aztec word ehecatl means wind, air,
life, soul, shadow. A common term in the Old Testament, nephesh
means living spirit or breath-soul, and to die is to "breathe out the
nephesh." The Chinese ch'i, irnportant both to martial arts and med-
icine, is the life force, the cosmic spirit that pervades and enlivens
all things and that is synonymous with primordial energy. Although
for the ancient Greeks psyche meant "breath-soul" (as distinct from
"blood-soul," which was the seat of consciousness), by Aristode's
time psyche had come to mean the life principle. Pneuma, another
Greek term for spirit or soul, familiar to us in "pneumonia" or "pneu-
matic," derives from the word pnein, which means to breathe. In De
Anima ("On the Soul") Aristotle claimed the soul, the purpose for
which the living body exists, is the source of motion ("animation").
The magico-religious notion of a holy spirit breathing souls into
bodies worked its way into science. Before the eighteenth century
live bein5 were not said to "reproduce"; they were "generated."
 36 What ls Life?

The monsters of bestiaries were the combinatorial results of souls

and natural and divine intervention. Aristotle thought that by way
of semen men contribute souls, whereas women provide the nur-
turing matter in the generation of the child. "Parents are merely
the seat of the forces uniting matter and form," wrote Jean Franqois
Fernel (r+gl-r558), who coined the terms physiology and pathology
and who was physician to King Henry II of France. 'Above them
stands a more powerful Workman. It is He who determines the form
by breathing the breath of 1ife."2
The observation that some things, such as rocks, lacked animat-
ing souls implied the obverse, that self-moving spirits floated bod-
ilessly through the ether. Combined with a longing for immortal-
iry this inference-that souls exist on their own-offered hope of
cheating death. The notion of disembodied spirit is at the root of
ancestor worship and beliefs in ghosts, angels, and reincarnation. For
Plato the heavens were inhabited by ensouled planets and stars, the
world a divine repetition, within time, of a perfect realm beyond
time: a universe of pure mind. In Timaeus he wrote, "The world
has received animals, mortal and immortal, and is fulfrlled with them,
and has become a visible animal containing the visible-the sensi-
ble God who is the image of the intellectual, the greatest, best, fairest,
most perfect-the one only-begotten heaven." Aristotle, who stud-
ied living beings well enough to correctly describe the use of ten-
tacles in mating octopi, modified Plato's ideas. He emphasized the
mundane purpose of living beings as the great purpose generated
by "First Cause" or "LJnmoved Mover." Christianiry influenced by
Greek philosophy through the Church fathers, incorporated the He-
brew notion of a single God. Christian doctrine dispensed with na-
ture, spirits, and ancillary gods save those, such as saints and angels,
who mediate berr,veen man, his soul, and God. The souls and spir-
its once perceived to be rampant throughout nature became van-
ishingly scarce.
In the Middle Ages (c. 5oo-r 5oo c.E.) a European religious sect
known as the Gnostics decided that the true self was a divine spark
 Lost Souls I l,

trapped in a prison of fleshly matter. The Gnostics pictured Earth

as surrounded by seven transparent, crystalline spheres-the clear
heavens-each of which held a celestial body: Moon, Mercury,
Venus, Mars, Jupiter, Saturn, and Sun. The spheres holding the plan-
ets were living spiritual powers, three-dimensional glass ceilings, so
to speak, presided over by Archons, cosmic bouncers whose busi-
ness it was to prevent souls from rising back to heaven. Gnosticism
made sense in a medieval Europe. The Black Plague-half-dead bod-
ies moaning and rotting in the streets, warmer ones flagellating one
another and predicting the end of the world-called into question
the behavior of a Creator much as the Holocaust has done in our
time. But whereas existential philosophers sided against God's ex-
istence, Gnostics argued only for His absence.
During the Renaissance, when the classical Greek and Roman
texts-which had been protected during the so-called Dark Ages by
Islanric scholars-were rediscovered, thinkers risked their lives to free
themselves from religious dogma. Giordano Bruno (r 548-r6oo) was
burned at the stake for his seven years of resolute heresy. Espousing
a pantheistic perspective in which God, life, and mind were part of
an ever-changing universe, Bruno even thought distant worlds might
harbor intelligent beings. The same Christian view that Bruno defied
holds firm today: God is as superior to the universe as mind is to
matter, or soul is to body. Flesh, a necessary evil, is unclean; only
spirit is pure.

CARTESIAN LICENSE

At the dawn of modern science, French Catholic mathematician

Ren6 Descartes (r596-165o) posited a fateful split berween res ex-

tensa, material realiry and res cogitans, thinking reality. Only humans,
Descartes argued, partake of God to the extent that they have souls.
Even animals, though they seem to Gel pain, are soulless machines:
"We are so accustomed to persuade ourselves that the brute beasts
feel as we do that it is difficult for us to rid ourselves of this opin-
 38 What ls Life?

ion. But if we were as accustomed to seeing automata which imi-

tate perfectly all those of our actions which they can imitate, and
to taking them for automata only, we should have no doubt at all
that the irrational animals are automata."3
On the authority of Descartes, live animals were nailed to boards
without remorse to illustrate facts of anatonry and physiology. Nev-
ertheless, Descartes's presentation of the universe as a vast mecha-
nism did serve to open up the cosmos for scientific investigation.
Unfeeling nature could be analyzed with no fear of trespass. Na-
ture, a vast lifeless mechanism, could be dismantled and rnanipu-
lated, experimented upon with impunity. Man became the final
earthly refuge of divine presence.
By splitting reality into human consciousness and an unfeeling,
objective, "extensive" world that could be measured mathematically,
Descartes paved the way for a scientific investigation of nature con-
structed according to the mathematical laws of God.
"God sets up laws in nature just as a king sets up laws in his king-
dom,"4 wrote Descartes. A kind of Cartesian license gave prece-
dence to matter over form, body over soul, ourward spatially ex-
tended nature over inner awareness. Matter, body, and nature
could-unlike thought or feeling-be quantified, examined, and
ultimately understood by mathematical physics.
This Cartesian license permitted the human intellect, through sci-
ence, to enter a thousand different realms, fiom the very small to
the very large, and even the invisible. The blueprint underlying the
great mechanism of the cosmos was thought decipherable. Optical
instruments were focused on snowflakes and peppercorns, or pointed
at the pockmarked whiteness of the side-lit moon. Atoms were in-
vestigated by chemical combination and physical acceleration. X-
rays imaged bones. Radioactive elements clocked the internal me-
tabolism of the human body. Engineers even appropriated the
seenringly God-given power to fly.
Flashing the Cartesian permit to practice science did yield results.
Investigators returned the Bible and the classic books to their dusty
 Lost Souls I Sg

shelves. Instead they read Nature, "written," as Galileo Galilei

(r 564-1642) had put it (even before Descartes), "in a great book
which is always open before our eyes."s Galileo paid dearly for his
scientific leanings. As quantitative mechanist, measurer of falling
bodies, discoverer of the moons of Jupiter and the rotation of the
sun, Galileo cleared the trail for curious successors. As defier of po-
tent philosophers and Christian theologians, Galileo provoked the
ire of Church authorities. Although he was not, like Bruno, burnt
at the stake, he was, at age 58, brought before the Inquisition and
charged with heresy. Galileo recanted his earlier claims so at vari-
ance with official Church doctrine and "admitted" that Earth is the
center of the universe. 'Warned against further heresy, he was con-
demned to three years of weekly psalm recitations. He became a
prisoner in his own country home. Galileo's imrnensely popular
masterpiece, Dialogue of the Two Chief World Systems, was banned-
until I 83 8. Pope UrbanVIII (r 568-r644), believing that he had been
mocked as spokesman of the Church's cosmological views in
Galileo's character "Simplicio," began the censorship.
If Galileo had worked under Cartesian permit he would have fared
better. The devout Descartes abandoned work on a book manuscript
in which he was putting forth similar views, when in I63 3 he learned
of Galileo's condemnation. Anxious to conciliate religion and sci-
ence, Descartes gave great impetus to modern philosophy by doubt-
ing everything but the existence of his own doubting mind. The
body was entirely mechanical, he held, but connected to the mind
through the pineal gland, a pea-sized structure known at the time
only in human brains. The pineal gland acted, Descartes believed,
as a valve through which God was connected to the human soul.
To this day the Cartesian permit rallies scientists to study a uni-
verse that is wide open for investigation, but in the "fine print" is
found the exception: the conscious human soul-which in Des-
cartes's time was unquestionably made in God's image. Moreover,
the Cartesian permit still contains in the fine print this assumption:
the universe is mechanical and set up according to immutable laws.
 40 What ls Life?

Neither the exception nor the assumption is science. At the very heart
of the Cartesian philosophy are thus metaphysical presuppositions,
springing from the culture that gave rise to science.
Ultimately-in our very abbreviated story-the Cartesian license
proves to be a kind of forgery. After three centuries of implicit re-
newal, the license is still accepted even though the fine print, erased
or ignored, is no longer visible at arry magnification. Yet this fine
print was not incidental. It was the raison d'6tre, the rational basis
authorizing scientists following the spirit of Descartes to proceed
with their work and to receive the blessings of sociery if not always
the Church. The Cartesian view of cosmos as machine is at the very
root of the practice of science.

ENTERINC THE FORBIDDEN REALM

-While
Descartes cogitated, Europe remained under the rule of roy-
alry. The King and the Lord, representing the power and order of
God, reigned supreme. But science soon entered the forbidden realm
of humankind, the one place it was not supposed to go. Scientific
revelation of mechanism, part of the new audaciry of inquiry, helped
unsettle European monarchy. If the universe made by God is a gi-
ant automaton that works itself, why should people obey any King
or Lord whose power, God-given in the feudal system of medieval
Christianiry no longer derived from heavenly decree? The high-born
Frenchman Donatien Alphonse Frangois de Sade Q74o-r8 r4) keenly
felt the vanishing basis for moraliry. If Nature was a self-perpetuating
machine and no longer a purveyor of divine authoriry then it did
not matter what he, as the infamous marquis de Sade, did or wrote.
In r776 the British colonists in North America broke free from
transatlantic rule. Independence from the burdens of taxes and roy-
alry was proclaimed. In r789 the French Revolution deposed the
king and stripped the lords and ladies of their powers. Irreverent
Voltaire Q694-r778) claimed that if God did not exist it would
be necessary to invent him. (A century later German philosopher
 LostSouls I O't

Friedrich Nietzsche [r 844-r9oo] would declare God dead.)England,

too, was struck by the revolutionary spirit of the time, but in mod-
eration. Retaining their king and queen, theEnglish perceived them-
of order in a world gone mad.
selves a bastion
Enter Charles Darwin. In r 859 his Origin of Species was published,
announcing to the world the scientifically derived inference that man
had not been created by God, but had evolved from mere animals
through "natural selection." Darwin's later books, Descent of Man
(r 87 r) and Expression of Emotions (t872), explored the then-startling
thesis that humans and apes evolved from ancient apes. Darwin
documented, without any explicit anti-Christian statement, that
neither humans nor ancestral apes were created by God. The Great
Chain of Being-the line of holiness coming down from God
through spiritual angels to humankind and thence to the rest of me-
chanical creation-was turned topsy-turvy. The cosmic apple cart
was upset. No longer, Darwin insinuated, was Man excluded from
connection with nature. Even the perceiving rnind, describing it-
self, evolved from mechanical laws of random variation and natu-
ral selection. Materialism was victorious. As in some maudlin Dis-
ney animation, the last sparkle of fairy stuffdisappeared.
'Western
thought thus suffered a metaphysical reversal. Once, be-
fore the exploits of Bruno and Galileo, Descartes and Newton, and
Darwin, everything had been alive, except for the natural magic trick
of death; now, in the scientific-mechanistic world, everything was
inanimate, dead, except for the scientific puzzle of life.
We all are interested in life because we know it from the inside
as something more than mechanical, automatic, deternrined re-
sponses to preordained stimuli. We think, act, choose. We-and it
would be a conceit to exclude other organic beings-are not New-
tonian machines.
Moreover, we are not objective outsiders. In physics, 'Werner
Heisenberg's uncertainry principle limits what is measurable. In
mathematics, Kurt Gcidel's incompleteness theorem warns that
every mathematical system, if complete, cannot be consistent and,
 42 What ls Life?

if consistent, cannot be complete, since to define it axioms are

needed from outside the system. Such scientific uncertainry also im-
pedes any search to define life. On the one hand, a final definition
of life by life may be like kissing your elbow or rolling your eyes to
see your own optic nerve: impossible. On the other hand, enlighr
ened by a knowledge of history and science's astounding success at
investigating what life is, we seem closer than ever to a deeper un-
derstanding of life in its cosmic and cultural context.
In the flush of this exhilarating material success scientists tend to
gloss over the distinction berween life and nonlife, pointing to the
chenrical continuities. Life-as-a-whole is like other vast subjects: na-
tionalism, culture, politics, or anything else not easily defined, ma-
nipulated, or described. Even biologists may be srude, dismissing rel-
evant discussion as 'just philosophy." But science, like anything else,
has a context. And that context is partly metaphysics, great, often-
unstated categories of thought, perhaps cultural, perhaps inherited
(the distinction is itself metaphysicall) that go beyond science
proper. No one escapes metaphysics; to understand life, as science,
it is necessary to understand its cultural context.
"Metaphysics," introduced by Hellenistic scholars and referring
to certain untitled texts by Aristotle, comes from the Greek ta meta
ta physika biblia, which literally means "the books after ('meta') the

books on nature." The original use of the prefix "meta," by early

editors such as Andronicus of Rhodes, may not have referred to any
transcendental interpretation of ultimate realiry but only to the
mundane position of the book on the table where "Metaphysics"
was stacked on top of "Physics." Beginning with the work of Im-
manuel Kant, metaphysics has come to reGr to speculations on ques-
tions not answerable by direct observation or experiment. Meta-
physics, as a web of ideas in which we are caught, need not give
rise to futiliry. It is fascinating to try and tease apart the strands of
the culturally inherited, linguistically reinforced concepts that guide
even our most seemingly original thoughts. An explanation of meta-
physics may not lead to absolute truth, but it certainly shouldn't be
anathema to open, scientific minds.
 Lost Souls I O,

cosMrc wrGGLEs

'A living body," wrote Alan Watts (r9r 5-t973), "is not a fixed thing
but a flowing event." 'W'atts, the Anglo-American popularizer of
Eastern philosophy, drew from science, as well, in his quest for the
meaning of life. He likened life to "a flame or a whirlpool":

The shape alone is stable. The substance is a stream of energy going

in at one end and out at the other. Life's purpose to maintain and
perpetuate itself is understandable as a physico-chemical phenome-
non studied by the science of thermodynamics. 'We are temporarily
identifiable wiggles in a stream that enters us in the form of light,
heat, air, water, milk. . . . It goes out as gas and excrement-also as
semen, babies, talk, politics, war, poetry and music.6

Thermodynamic systems lose heat to the universe as they convert

energy from one form to another. Living matter frees itself from or-
dinary matter only by perpetually basking in the sun. Confronted
with dissolution and destruction, life suffers a permanent death
threat. Life is not merely matter, but matter energized, matter or-
ganized, matter with a glorious and peculiar built-in history. Life as
matter with needs inseparable from its history must maintain and
perpetuate itself, swim or sink.The most glorious organic being may
indeed be nothing but "temporarily identifiable wiggles," but for
millions of years as life has been racing away &om disorder, au-
topoietic beings have concerned themselves with themselves, be-
coming ever more sensitive, ever more future oriented, and ever
more focused on what might bring harm to the delicate wave of
their matter-surfing fornr. From a thermodynamic, autopoietic per-
spective, the basest act of reproduction and the most elegant aes-
thetic appreciation derive from a common source and ultimately
serve the same purpose: to preserve vivified matter in the face of
adversity and a universal tendency toward disorder.
Dutch-Jewish philosopher Baruch Spinoza (t 6 3 z-r 67 7) portrayed
matter and energy as the fundamental nature of a universe which
was itself alive. The great German writer and naturalistJohann Wolf-
gang von Goethe $749-l.832), author of Faust, argued for a po-
 44 What ls Life?

etic biology. He thought matter does not operate without spirit, nor
does spirit existwithout matter. Although he was pre-Dar-winian
and his theories are now obsolete, Goethe wrote ably on science.
In one passage he plucks from human activiry what mtght be called
its autopoietic essence:

'W'hy are
the people thus busily nroving? For
food they are seeking,
Children they fain would beget, feeding them
well as they can.
Traveler, mark this well, and, when thou art
home, do thou likewise!
More can no mortal effect, work with what
ardor he wi11.7

The German biologist Ernst Haeckel (r 8 3 4- r 9 r 9), inventor of the

word "ecology," promoted the idea that the activiry of the human
psyche is an offshoot of physiology: "We hold with Goethe that
matter cannot exist . . . without spirit. . . . We adhere firmly to the
pure, unequivocal monism of Spinoza: Matter, or infinitely extended
substance, and Spirit (orEnergy), or sensitive and thinking substance,
are the two fundamental attributes, or principal properties, of the
all-embracing essence of the world, the universal substance."s

THE MEANING OF EVOLUTION

Ernst Haeckel was Darwin's translator and greatest advocate in the

German tongue, but he pushed Darwinism further than its inven-
tor had been willing to carry it. The soul, Haeckel claimed, resided
in the cell, immortaliry was a metaphysical sham, life had no pur-
pose other than itself, and beings were not spiritual but material in
nature. "FIumaniry" he declared, "is but a transitory phase of evo-
lution of an eternal substance, a particular phenomenal form of mat-
ter and energy, the true proportion of which we soon perceive when
we set it on the background of infinite space and eternal time."e
 Lost Souls I OU

Such views infuriated traditional religious sensibilities, including

those of Alfred Russel Wallace (r 823-1 9 r 3 ). An English naturalist,
'Wallace
developed his own theory of evolution by natural selection
that was uncannily similar to that of Darwin. Darwin's and'Wal-
lace's short papers on natural selection were published together in
the same issue of the Journal of the Proceedings of the Linnean Soci-
ety oJ London, Zoology. Wallace, who frequented seances, reviled
Haeckel's notion of matter as eternal and aLive, and he rejected
Haeckel's denial of a spirit world. He sneered that the riddle of the
universe-which was the title of one of Haeckel's most influential
and popular books-had not been solved, least of all by Haeckel.
Even before Darwin, German philosopher Immanuel Kant (r724-
r8o4) noted that skeletal and other similarities pointed to blood
ties, a common parentage for all life. Kant ceded that all life could
have arisen through some mechanical process similar to that by
which nature produces crystals, but he judged it would be absurd
to hope for "a Newton" who could make comprehensible even the
growth of a single blade of grass by mechanical theory alone.
Haeckel proposed Darwin as the very "Newton" Kant had believed
impossible.
By projecting Earth history millions of years beyond the six
thousand years allotted in the Book of Genesis, James Flutton (r726-
ry97) founded modern geology. Hutton, son of a Scottish merchant,
distinguished rocks laid down as sediment from those brought forth
in molten form through volcanoes. He observed erosion by wind
and water and deduced the production of rainfall from cooling air
masses that could no longer contain their moisture. Older sedi-
ments were deposited prior to more recent ones. Hutton's "law of,
superposition" led to Charles Lyell's (r799-r87 5) statement of the
"law of uniformitarianism," the suggestion that only those geolog-
ical forces observable in the present need be invoked to account for
structures made and sediments accumulated in the past. But Hut-
ton's extrapolation that Earth must be very old was controversial.
Conservative England, threatened by the wild and godless French
 46 What ls Life?

Revolution, was not ready to accept an Earth older than that which
could be ascertained by sumrning up all the "begats" mentioned in
the Bible.
Nonetheless, Scottish geologist Charles Lyell approved Hutton and
argued that time was far vaster than previously thought in his mul-
tivolume book, The Principles of Ceologlnwhich did for that field
what Darwin's opus later did for zoology and botany. Lyell was also
far ahead of his time in taking a global ecological perspective rem-
iniscent of Gaia theory today; he called attention to "the powers of
vitality on the state of the earth's surface."10 Darwin read Lyell dur-
ing his voyage on the Beagle and adopted the Lyellian worldview.
Decades laterLyell, in turn, embraced the Darwinian worldview. In
r863 he published The Antiquity of Man, which suggested, before
Darwin had made the extension, that evolution applied to all hu-
mankind.
Meanwhile on the Continent, Berlin naturalist Christian Gottfried
Ehrenberg (rlgS-t876) was putting the life back into biology. Re-
turning from an ill-fated expedition to Egypt, of which he was per-
haps the sole survivor, he focused on the transition between life and
nonlife. In the expedition to Egypt (r 8zo) and a later one to Siberia
(r829) Ehrenberg documented the unseen world of nricrobes that
fertilize the oceans and soils. Through his journeys Ehrenberg came
to know Friedrich Wilhelm Alexander von Humboldt (r 769- I 8 5 9).
The baron von Humboldt, widely regarded as the greatest German
naturalist of his time, had collected more than sixry thousand plant
specimens during his travels around the world. He had visited Amer-
ican president Thomas Jefferson (r74-18z6) and was described as

a scientific"Napoleon." In his seventies Humboldt began to com-

p17e Kosmos, his grand attempt to map and explain the entire uni-

verse. "Certainly," wrote Isaac Asimov (r9zo-rg9z), "no man be-

fore him, with so active a rnind, had seen so much of the world, and
no man before him was so well equipped to write such a book. . . .
It was a florid production, rather overblown, but it is one of the
remarkable books in scientific history and was the first reasonably
accurate encyclopedia of geography and geology."rl
 Lost Souls I O,

In Humboldt sharesEhrenberg's discovery of liG's global

Kosmos,
sweep. "The universality of life is so profusely distributed," waxes
Humboldt,

that the smaller Infusoria [ciliates and other protists] live as parasites
on the larger, and are themselves inhabited by others. . . . The strong
and beneficial influence exercised on the feeling of mankind by the
consideration of the diffusion of life throughout the realms of nature
is common to every zone, but the impression thus produced is most
powerful in the equatorial regions, in the land of palms, bamboos,
and arborescent ferns, where the ground rises from the shore of seas
rich in mollusca and corals to the lirnits of perpetual snow. The local
distribution of plants embraces almost all heights and depths. Organic
forms not only descend into the interior of the earth, where the in-
dustry of the miner has laid open extensive excavations and sprung
deep shafts, but I have also found snow-white stalactitic columns en-
circled by the delicate web of an Usnea [old man's beard lichen], in
caves where meteoric water could alone penetrate through fissures. . . .

[Organisms flourish on the summits of the] Andes, at an elevation

of more than r 5,ooo feet. Thermal springs contain small insects (Hy-
[iron bacterial masses], Oscillatoria, and
droporus thermalis), Callionellae
ConJeruae [an old name for a miscellany of green algae], while their
waters bathe the root-fibers of phanerogamic [cone- and flower-
bearing] plants.12

Humboldt died the same year Darwin publish ed The Origin of Species,
IJntil very recently, with publication of the work of Schrcidinger's
legacy, observations made by Humboldt and Ehrenberg on the mi-
crobial world and many other late nineteenth-century discoveries
were not brought together in an evolutionary context. The fertil-
ization of sperm by .gg (embryo formation), inheritance factors of
garden peas (Mendelian genetics), mucoid substances in the pus of
soldier's wounds (nucleic acids, DNA and RNA), and visualization
of chromosomes were some of the revelations made last century
which, in geneticist Theodosius Dobzhansky's words, only "make
sense . . . in the light of evolution."l3
Although theories of evolution had been in the air for a half cen-
tury and more, Darwin's methodical purposefulness, his diplomacy
 48 What ls Life?

of prose, and his presentation as an Englishman of a mechanical the-

ory during a time when Isaac Newton's theory of graviry was the
last word in science all helped make the appearance of his book an
epic event. As one woman of sociery wryly remarked on hearing
the news of her less-than-noble apish origins, "Let us hope it is not
true. But if it is, let us hope it does not become generally known."
Since T/re Origin o;f Species, the idea of evolution has become in-
creasingly accepted-overwhelmingly by scientists and respectably
by the public (particularly the educated public). But it has also been
abused. For example, in a popular illustration Haeckel depicted the
sumrnit of evolution as a nude but demure Germanic woman at the
top of his evolutionary tree. Haeckel's error was not so much in his
Germanic bias (or his choice of the female sex) but in his choice of
any human at all. This is because all extant species are equally evolved.
A11 living beings, from bacterial speck to congressional comnrittee
member, evolved from the ancient common ancestor which evolved
autopoiesis and thus became the first living cell. The fact of survival
itself proves "superioriry" as all are descended from the same me-
tabolizing [Jr-form. The gentle Living explosion, in a circuitous
4,ooo-mi1lion-year path to the present, has produced us all. In a sense
then, the Vedic intuition that individual awareness is illusory and
that each of us belongs to a single primal ground-Brahman-may
be accurate: we share a common heritage, not only of chemistry
but of consciousness, of the need to survive in a cosmos whose mat-
ter we share but which is itself indifferent to our living and self-
concern.

VERNADSKY'S BIOSPHERE

Given the limited legacy of metaphysical dualism (mind/body,

spirit/matter, lifelnonlife), it may not be surprising that fwo of the
most profound rethinkers of life and its environment in this cen-
tury share a biospheric perspective yet have diametrically opposed
views. But whereas Russian scientist Vladimir Ivanovich Vernadsky
 LostSouls I O,

(r863-1945) described organisms as he would minerals-calling

them "living matter"-English scientist James E. Lovelock de-
scribes Earth's surface, including rocks and air, as alive.
Vernadsky portrayed living matter as a geological force-indeed,
the greatest of all geological forces. Life moves and transforms mat-
as flying phosphorus-rich gulls,
ter across oceans and conrinents. Life,
racing schools of mackerel, and sediment-churning polychaete
worn$, moves and chemically transforms the planet's surface. More-
over, life is now known to be largely responsible for the unusual char-
acter of Earth's oxygen-rich and carbon dioxide-poor atmosphere.
Like Ehrenberg and Humboldt before him, Vernadsky showed
what he called the "ubiquity of life"-living matter's almost total
penetration into, and consequent involvement in, seemingly inan-
imate processes of rock, water, and wind. Others spoke of an ani-
nral, vegetable, and mineral kingdom; Vernadsky analyzed geologi-
cal phenomena without preconceived notions of what was and was
not alive. Perceiving life not as life but as "living matter," he was
free to broaden its study beyond that of biology or any other tradi-
tional discipline. What struck him most was that the material of
Earth's crust has been packaged into myriad moving beings whose
reproduction and growth build and break down matter on a global
scale. People, for example, redistribute and concentrate oxygen, hy-
drogen, nitrogen, carbon, sulfur, phosphorus, and other elements
of Earth's crust into rwo-legged, upright forms that have an arrraz-
ing propensity to wander across, dig into, and in countless other ways
'We
alter Earth's surface. are walking, talking minerals.
Vernadsky contrasted graviry which pulls matter vertically toward
the center of Earth, with liG-growing, running, swimming, and
flying. Life, challenging graviry. nloves matter horizontally across the
surface. Vernadsky detailed the structure and distribution of alumr-
nosilicates in Earth's crust and was the first to recognize the im-
portance to geological change of heat released from radioactiviry.
But even a resolute materialist like Vernadsky found a place for
rnind. In Vernadsky's view a special thinking layer of organized mat-
 50 What ls Life?

ter growing and changingEarth's surface is associated with humans

and technology. To describe it, he adopted rhe term noosphere, from
Greek noos, mind. The term had been coined by Edouard Le Roy,
philosopher Henri Bergson's successor at the Colldge de France.
Vernadsky and Le Roy met in Paris for intellectual discussions in
the r9zos, along with PierreTeilhard de Chardin (r88r-r955), the
French paleontologist and Jesuit priest whose writings would later
bring the idea of noosphere-a conscious layer of life-to a wide
audience. Teilhard's andVernadsky's use of the term noosphere, like
their slants on evolution in general, differed. For Teilhard the noo-
sphere was the "human" planetary layer forming "outside and above
the biosphere," while for Vernadsky the noosphere referred to hu-
maniry and technology as an integral part of the planetary biosphere.
Vernadsky distinguished himself from other theorizers by his
staunch refusal to erect a special category for life. In retrospect we
can see the value of his stance; because life has indeed become a
category, theorists of liG have managed to reifir-to make a thing
out of-something that is not a thing at all. Vernadsky's referring
to life as "living matter" was no mere rhetorical ploy. In one deft
verbal stroke Vernadsky cut loose centuries of mystic clutter attached
to the word "life." FIe made every attenlpt to consider life part of
other physical processes and consistently used the gerund "living"
to stress that life was less a thing and more a happening, a process.
Organisms forVernadsky are special, distributed forms of the com-
mon mineral, water. Animated water, life in all its wetness, displays
a power of movement exceeding that of limestone, silicate, and even

air. It shapes Earth's surface. Emphasizing the continuiry of watery

life and rocks, such as that evident in coal or fossil limestone reefs,
Vernadsky noted how these apparently inert strata are "traces ofby-
gone biospheres."l4
Austrian geologist Edward Suess (r 8 3 r-r 9 r 4) had coined the word
"biosphere," but VernadsLy brought it into currency. Just as the sphere
of rock is a lithosphere, and that of air an atmosphere, so the sphere
where life exists is a "biosphere." In his r9z6 book, The Biosphere,
 Lost Souls I U,

Vernadsky showed how Earth's surface was an ordered transforma-

tion of the energies of the sun. "The biosphere," wrote Vernadsky,
"is at least as much a creation of the sun as a result of terrestrial pro-
cesses. Ancient religious intuitions that considered terrestrial crea-
tures, especially man, to be children of the sun were far nearer the
truth than is thought by those who see earthly beings simply as
ephemeral creations arising from blind and accidental interplay of
matter and forces. . . . Living matter as a whole . . . is therefore a
unique system, which accumulates chemical free energy in the bio-
sphere by the transformation of solar radiation."ls
Remarkably, Vernadsky dismantled the rigid boundary befween
living organisms and a nonliving environment, depicting life glob-
ally before a single satellite had returned photographs of Earth from
orbit. Indeed, Vernadsky did for space what Darwin had done for
time: as Darwin showed all life descended from a remote ancestor,
so Vernadsky showed all life inhabited a materially unified place, the
biosphere. Life was a single entiry transforming to earthly matter
the cosmic energies of the sun (see plate l). Vernadsky portrayed
life as a global phenomenon in which the sun's energy was trans-
formed. Emphasizing photosynthetic growth of red and green bac-
teria, algae, and plants, he saw these expressions of living matter as
the "green fire" whose expansion, fed by the sun, pressured other
beings into becoming more complex and more dispersed.
Vernadsky set forth rwo laws. Over time, he claimed, more and
more chemical elements became involved in the cycles of life. Sec-
ond, the rate of migration of atoms in the environment has increased
with time. A flock of mrgrating geese was to Vernadsky a biospheric
transport system for nitrogen. Locust swarms, recorded in the Bible,
attested to massive changes in the distribution of carbon, phospho-
rus, sulfur, and other biologically important chemicals fwo thou-
sand years ago. As dams, factories, mines, machine construction,
utilities, trains, planes, global communications, and entertainment
systems have appeared, more chernical elements than ever have be-
come organized into functioning parts of autopoietic systems. Tech-
 52 What ls Life?

nology, from a Vernadskian perspective, is very rluch a part of na-

ture. The former calf muscle severed into brochette cubes and the
pine tree trunk into lumber pass through the hands of workers and
the chutes of machines to emerge transformed into shish kebab and
flooring. The plastics and metals incorporated in industry belong to
an ancient process of life co-opting new materials for a surface ge-
ological flow that becomes ever nlore rapid. And, with the fleeting
synthesis in physicists' laboratories of radioactive isotopes, the noos-
phere begins to direct and organize atoms that have never before
existed on Earth.

LOVELOCK'S GAIA

As Vernadsky disrupted the mind/marter split through a consider-

ation of living matter on a global scale, so James E. Lovelock up-
sets metaphysical dualism by an opposite stratagem-considering
Earth alive. Vernadsky examined life as matter within a receptive
political and cultural climate-the official atheism of the former
Soviet lJnion, aided by science's approval of materialism. By con-
trast, Lovelock-portraying the self-regulating biosphere, a huge and
oddly spherical living body he calls "Gaia"-has been hampered by
the subtle ideology of mechanism that pervades the scientific com-
muniry. This means that Lovelock must not only show that Earth
maintains itself as a living body, he must also overcome the preju-
dice that to call this "thing" alive is not science but poetic per-
sonification. Given these tensions, it is a restament to this world-
class atmospheric chemist's ingenuity that his theory is taken as
seriously as it is by active scientists.l6
Atmospheric, astronomical, and oceanographic evidence attest
that life manifests itself on a planetary scale. The steadiness of mean
planetary temperature for the past 3,ooo million years, the 7oo-
million-year maintenance of Earth's reactive atmosphere befween
high-oxygen levels of combustibility and low-oxygen levels of as-
phyxiation, and the apparently continuous removal of hazardous salts
 Lost Souls lu,

FIcURE 5. Emilianahuxleyi, a
coccolithophorid. Phylum: Hap-
tomonada. Kingdom: Protoc-
tista. This coccolithophorid, a
calcium-precipitating alga, is
covered with button-like scales.
These protists, each only 20 mil-
lionths of a meter in diameter,
produce dimethyl sulfide, a gas
of global significance involved
in cloud cover over the ocean.

from oceans-all these point to mammal-like purposefulness in the

organization of life as a whole (fig. S).
This purposefulness, central to scientific Gaia theory, is a major
sticking point for traditional biologists. How can a planet behave
in a purposeful manner to maintain environmental conditions fa-
vorable to its living constituents? In mechanistic biology, complex
self-regulation only evolves from natural selection that weeds out
more poorly self-regulating individuals. This logic is flawed, how-
ever. According to it no original, self-maintaining cell could have
ever evolved, because "purposeful," self-regulating behavior simply
cannot arise in a population with only one member. A strict read-
ing of Darwinism
G
?
,
Plausible within the bounds of Darwinism or not, both planet,
isolated by space, and cell, isolated by semipermeable membrane,
are solar energy-requiring systems, continuous through time and
space, that display self-maintaining behavior. The "purposefulness"
of Gaian self-maintenance derives from the living behavior of myr-
iad organisms, mostly rnicrobes, whose ubiquity Ehrenberg and
Humboldt first established. Planetary physiology, far from having
 54 | wf,"t ts Life?

been produc ed ex nihilo, or by an outside God, is the holarchic out-

come of ordinary living beings. It is the autopoiesis of the cell writ
large.
Life cannot be understood while ignoring the sentient observer.
If not for mind, no one would care thar life is a certain kind of sun-
light-energized cosrnic debris. But it is, and we do. To best under-
stand life we need to see the long and winding road from animism,
through dualism, to the limrtations of mechanism. Physics, chem-
istry, and biology are distinct approaches to the same material phe-
nomena. As German geornicrobiologist Wolfgang Krumbein puts it,

The mineral and rnicrobial mineral cycles as we view them today on

the basis of experimental work have been envisaged as the uniflzing
concept of world and universe, creating the principle of the one liv-
ing nature of Bruno and Spinoza. . . .The basic approach of Bruno . . .
is still alive and is evidenced in scientific and mathematic terms by
non-Euclidian geometry, by the modern field theories and Einstein's
relativiry and gravity theories, as well as by Lovelock's "Gaia-
hypothesis." Giordano Bruno deeply influencedBaruch Spinoza fifiz-
r 677], Leibnitz 1t646-r7 t6l, Kant j7
z4-r8o4), Goethe 17 49-r 8 3zl
and Schelling 11775-r854]. He still influences unirarian thoughr in
science andphilosophy . . . a textbook on microbial geochemistry . . .
must come back to Bruno's original thoughts of "cyclic develop-
ments" rather than "creation and destiny" as revealed in the clerical
Christian thoughts of his time which have so severely inhibited the
development of science. 17

Life can be returned to biology without compromising science.

Mechanism gave science the authoriry to examine the realms of,
heaven and life once considered "offlimits." But it also suggested
the universe was more deterministic than it is, cutting into our sense
of life and wonder. The Epicurean Roman philosopher Lucretius
(SS-SS B.c.E.), in his poem De Rerum lrlatura ("On the Nature of
Things"), presents an evolutionary view of the universe denying a
hereafter and arguing that everything, even the soul and gods, is
made of atoms. In the same tradition, Bruno blended matter with
 Lost souls I UU

energy, finite with infinite, world with God. In the modern era, by
not speaking of life at all-but calling it "living matter"-Vernadsky
offered us a chance to see life with fresh eyes. And, unlike mono-
lithic Cartesian materialism, the Gaia perspective accommodates the
enchantment we feel as living beings dwelling in a living world.

SO, WHAT lS LIFE? Life is planetary exuberance, a solar phenom-

enon. It is the astronomically local transmutation of Earth's air, wa-
ter, and sun into cells. It is an intricate pattern of growth and death,
dispatch and retrenchment, transformation and decay. Life is the sin-
gle expanding orgarization connected through Darwinian time to
the first bacteria and through Vernadskian space to all citizens of the
biosphere. Life God and music and carbon and energy is a whirling
as

nexus of growing, fusing, and dying beings. It is matter gone wild,

capable of choosing its own direction in order to indefinitely fore-
stall the inevitable moment of thermodynamic equilibrium-death.
Life is also a question the universe poses to itself in the form of a
human being.
'V/hat happened to living matter to make it so different? The an-

swer is both scientific and historical. Life is its own inimitable his-
tory. From an everyday, uncontentious perspective, "you" began in
your mother's womb some nine months before whatever your age
is. From a deeper, evolutionary perspective, however, "yot" began
with life's daring genesis-itssecession, more than 4,ooo million
years ago, from the witches' brew of the early Earth. In the next
chapter we see how this brew, sometimes called the primeval soup,
started percolating.
 ONCE UPON A PLANET

lf a dirty undergarment is squeezed . . . a ferment drained

from the garments and transformed by the smell of the grain,
encrusts the wheat itself with its own skin and turns it into
mice. . . . And, what is more remarkable, the mice from corn
and undergarments are neither weanlings or sucklings nor
premature but they jump out fully formed.

JEAN BAPTISTE VAN HELMONT

For the humblest organism, the simplest bacterium, is already

a coalition of enormous numbers of molecules. lt is out of the
question for all the pieces to have been formed independently
in the primeval ocean, to meet by chance one fine day, and
suddenly arrange themselves in such a complex system.
FRANeOTS JACOB

It must be admitted from the beginning that we do not know

how life began.
STANLEY MILLER AND LESLIE ORGEL

BEGINNINGS

On Earth some 4,ooo million years ago life generated as matter un-
dertook a different direction. From the beginning, life satisfied its
autopoietic imperative in a universe obeying thermodynamic laws.
Bound and separated from the world by a border of its own mak-
ing, life came together as oily droplets that increased their order (see
plate 4). Other dissipative systems in nature use energy to increase

57
 What ls Life?

order, but they last for only short periods of time. Moreover, a tor-
nado risen on the plains doesn't go "whoops" as it wanders into a
mountainous landscape that spells its doom; but even the simplest
life form effectively does, actively responding to irs surroundings to
preserve and protect itself.
How matter in a bath of energy (or how energy in a brew of mat-
ter) first accomplished the feat of life is nor known. No molecule
alone can reproduce. Minimal life on Earth today is a system, a
minute membrane-bounded sphere, a bacterial cell, requiring many
interacting molecules. Some 5oo to j,ooo genes make a sirnilar num-
ber of proteins. Proteins and DNA mutually produce each other
within the cell membrane that together they fabricate. Bearing a
corunon biochemistry, all life probably dates to a single, perhaps
(but not necessarily) improbable historical moment. The factors thar
led matter to its peculiar "fractionation point" where dissipative be-
havior became living behavior need only have happened once. En-
closed, perhaps even suddenly, by a membrane and with resources
aplenry the first living cells could afford ro be somewhat aloof from
external realiry. Eventually, imperiled by its own profligacy and by
the insensitivity of the substance from which ig 5s6sclsd-yet upon
which it absolutely depended for sustenance-life was left to its own
devices. As matter ostracized from itself, life had been abandoned
by the world, yet the world had gone nowhere. There was no go-
ing back.
Once begun, reproducing systems proceeded rapidly away from
their initial state, and today no vestiges of ear\ life less complex
than a bacterial cell remain. Bacteria are not half-hewn but fully liv-
ing and evolved beings that have been thriving for more than 3,5oo
million years. The greatest chemical inventors in the history of Earth,
they are not 'just germs." Because of the conservative material na-
ture of reproducing life, bacterial cells retain clues to the chemistry
of Earth's surface it existed in the remote past. Bacteria were the
as
first green beings to grow on nothing but sun, water, and air. Still
the only beings able to perform many metabolic tricks of which we
 Once Upon a Planet I uS

animals and even plants are not capable, bacteria were the first to
breathe oxygen and to swim. They are the virtuosi of the biosphere.
They are also our relatives, which probably explains why we feel
free to malign them.
Bacteria, never having gone extinct, continue to protect us as
their populations grow prodigiously. They maintain soils for us and
purify waters. Bacteria expel gases, filling their immediate envi-
ronment with wastes noxious to the producers but alluring to other
strains as they colonize ubiquitous niches-even glacial ice and
boiling hot springs. Some build durable structures and crowd them
with their communities; some make vinegar; others work metals
such as iron, manganese, and even gold. Some sense the sun, swim-
ming to bask in its rays, while others are photophobic. Still other
bacteria sense and swim toward the closest magnetic pole. Many
bacteria are poisoned by oxygen; others thrive in it. Some make
spores remarkably resistant to heat, desiccation, or radiation. Bac-
teria come in a variery of colors, from snow-white Beggiatoa and
yellow sulfur bacteria to red Chromatium and blue-green cyanos
like Spirulina, Nostoc, or Microcystis. Bacteria, in short, are no more
"germs" than the plants that feed, clothe, and house us are
"weeds."
But how did the first bacterium originate? Again, no one knows.
Bacteria are so sophisticated that they could have come from space.
In the fifth century B.c.E. the Greek scientist Anaxagoras, friend of
the plal.wright Euripides, invented "panspermia," the notion that
life, dispersed as seeds throughout the universe, landed on Earth.
Later, Swedish chemist Svant6 Arrhenius ( I 8 59-r 9zg), Nobel Prize
winner for his ion theory that atoms in solution generate electri-
cal charge, proposed that hardy bacterial spores were pushed by
solar winds from star to star. Arguing that minute particles would
rise to the upper atmosphere in volcanic eruptions, and that some
of these particles, carrying bacterial spores, would reach the
stratosphere-where electrical discharges would propel them into
space-Arrhenius calculated that an Earth spore so launched today
 60 | wh"t ls Life?
would reach Pluto in four months and arrive at Alpha Centauri,
the nearest star, injust seven thousand years. (Spores discovered in
peat and other deposits older than that have proved to be viable.)
More recently, Francis Crick, codiscoverer of the structure of
DNA, argues for "directed panspermra"-that intelligent extra-
terrestrials may have seeded Earth with the cosmic equivalent of a
start-up kit for life.1
Might life have begun in some other star system and then have
migrated (or been exported) to Earth? Perhaps, but such a view is
less amenable to scientific investigation than the view that lifb orig-
inated right here on Earth. Moreover, if life started in outer space-
say, on an Earthlike planet-study of how life arose would apply to
its beginning anywhere. Indeed, Earth itself is suspended in space,
so any way we look at it, life came from space.

HELL ON EARTH

It is dawn, 4,6oo million years ago. Earth is in the violent red throes
of its beginnings, a gravitational implosion of molten rock and
swirling metal. Superheated gases such as ammonia, hydrogen
sulfide, and methane curl in an atmosphere zapped everl,where and
at every moment by lightning. The very oceans hang unfallen,
unrained-a sphere of steam eclipsing any semblance of sun. Be-
neath these vapors, thick with formaldehyde and cyanide (simple
organic compounds that form naturally in space), the boiling crust
of the surface pullulates with radioactivity and heat.
Meanwhile, the sun has ignited, flashing with a burst of radiation
so forceful it blows offwhole planetary atmospheres, relegating hy-
drogen gas to the outer reaches of the solar system. There hydro-
gen collects around cold and massive Jupiter, Saturn, (Jranus, and
Neptune-for only these giant planets have graviry suflicient ro re-
tain their original allotment of this lightest of all elements. Every-
where, on all planets and their moons, meteors ranging in size from
dust specks to large planetoids continue their bombardment. Smash-
 Once Upon a Planet 61

ing their way around the solar system, such space rubble brings wa-
ter and carbon compounds along for the ride, thickening the brew
that will feed early life on Earth.
One particularly huge intruder strikes Earth, flinging continent-
sized masses into space. But it is slowed enough by the collision to
become entrapped in Earth's orbit. After receiving a few more
smaller smashes of its own, the cratered orb comes to resemble the
white, sun-glinting moon that enchants us today. But in that far-off
era the scene would not have been serene. The young Earth spun
so fast that daylight lasted a mere five hours. The atmosphere, lack-
ing oxygen, would have supplied no breath and no blue vistas.
Such was Earth as it might have appeared in the Hadean eon
4,6oo-4,ooo million years ago. Life may well have been around,
however, by the late Hadean-once the molten surface of Earth
had sufiiciently cooled and no extraterrestrial impact was large
enough to rearrange the entire crust at a single strike. This earli-
est violent period of Earth history, named after Hades, Greek hell
and abode of the dead, is the first of four long eons (see timeline,
pages 6z-8o).
Fossils, whether stony tree trunks, traces of worm burrows, foot-
prints along petrified shorelines, spores buried in lake sediments,
or oily chemicals from decaying leaves, are evidence of past life.
No fossils-not even a volcanic rock-survives &omEarth's Hadean
eon. The Hadean on Earth can only be inferred from measure-
ment of far older natter from meteors and from the moon. Some
of the oldest rocks dating from the following eon have, however,
survived, and these, indeed, define the onset of the Archean (4,ooo-
z,5oo million years ago). A few "unmetamorphosed" Archean
rocks-those not subjected to altering heat and pressure-retain
traces of life. Australian rocks 3,485 million years old contain
eleven or more types of recognizable fossil bacteria. The oldest
rocks on Earth today thus contain vestiges of life. No one knows
when life began, but liG is at least as old as we could empirically
know it to be.
EARTH HISTORY TIMELINES
Here our "human-centered" or distorted-scale timeline represents major groups of
organisms and events in Earth history as people and books usually porrray them: with
no regard for chronological symmetry in a world consisting only of large animals and
plants. 'We follow this with a true-scale timeline (beginning on page 64), which de-
nies center stage to people and other mammals and does not short shrift the first 4,ooo
million years of our planet's history. While it is often thought that nothing of inter-
est occurred on Earth until the origin of the skeletalized Cambrian marine animals
some i4I million years ago, a true appreciation of history requires that the great early
chapters of life's story not be dismissed. The true-scale timeline depicts some of the
most significant events in the story of life before the evolution in the Phanerozoic
Eon of familiar life forms.

HUMAN-CENTERED OR DISTORTED-SCALE TIMELINE

(Millionsof l

years ago)

0.005 - Appearance of cities

0.0'l - Holocene epoch
Appearance of agricultural urban centers based on agricultural cultivation
of grasses (wheat, rice, etc.)
0.05 : Appearance of culturally modern, cave-painting humans
0.20 - Appearance of anatomically modern humans (Homo sapiens)
1.65 - Quaternary Period
Pleistocene epoch
Homo erectus and Homo neanderthalensls in the Middle East, Africa,
and Europe
4 -:- Appearance of human ancestors (Australopithecus afarensis)
5 : Pliocene epoch
, Meditenanean basin entirely dry and major faunal "turnover event"
,r in Africa
I Diversification of Miocene apes (Proconsul) including hominids
g., Ram ap ithecus, australopithecines)
(e.

23 : Neogene Period
' Miocene epoch
First widespread grasslands

35 - Oligocene epoch
40 - Appearance of angiosperm herbs and trees with fruit
57 - Eocene epoch
55 - CENOZOTC ERA
Paleogene Period
Paleocene epoch
Beginning of mammal diversification, including spread of primate order
Abundant and widely dispersed mammalian fossils belonging to extinct
families and genera
Second largest extinction in history of life (including non-avian dinosaurs)
 100 - Opening of what will become the Atlantic Ocean
Appearance of flowering plants (angiosperms) and primates
(plesiadapiforms)
145 - Cretaceous Period
20O - Appearance of hard silica tests of diatoms
2O8 - Jurassic Period
245 - MESOZOTC ERA
Triassic Period
Largest extinction event in history of life
Beginning of breakup of Pangea continent
290 - Permian Period
Formation of extensive salt deposits, indlcating inland seas and coral reefs
and possible biospheric control of ocean salinity
Appearance of large amphibians; mammal-like reptiles; bird-like reptiles,
including dinosaurs; and shrew-like mammals
. Radiolarians and other protists abound in marine sediments.
300 - Extension of reef-building coral anlmals (coelenterates) and coralline
(rhodophyte) algae
323 - Pennsylvanian Period
Widespread large trees in swamps lead to coal forests
362 - Mississippian Period
Widespread occurrence of fish and amphibian vertebrates in fossil record
408 - Devonian Period
Appearance of armored fish and invertebrate marine animals
Land extensively covered by first forests
First appearance of plants with seeds
440 - Silurian Period
Appearance of terrestrial plants, rhyniophytes, with fungi in their roots
Beginning of widespread life on land
500 - Colonization of land surfaces by algae and insects
510 - Ordovician Period
Appearance of first (jawless) fishes
541 _ PHANEROZOIC EON
PALEOZOIC ERA
Cambrian Period
Appearance in fossil record of Cambrian hard-bodied animals (such as
trilobites) and "plants": foraminifera, dinomastigotes, radiolarians, and
red algae
4,600 _ HADEAN EON
Origin of Earth-Moon system and other solar system planets
 What ls Life?

SPONTANEOUS GENERATION

In Greek myths, goddesses issue from sea shells and mortals can be
turned into animals or trees. "In all the things of Nature there is
something marvelous,"2 wrote Aristotle, who, because he looked
to the real world rather than Greek myths for knowledge, is recog-
nized as the first biologist or naturalist of the'Western world. Nev-
ertheless, Aristotle accepted as fact the (to u$ mythlike notion that
matter suddenly springs to life.
'W'e
now think organisms reproduce, but our ancestors imagined
that life, through some sort of fathering principle, spontaneously
generated: God produced Eve from Adam's rib, meat decayed into
maggots, one thing became another. A certain perceptual logic of
profmity and likeness suggested that decomposing vegetation brings
forth insects and that fireflies could issue, as Aristotle taught, from
glinting morning dew. Augustine (l+S-+:o) argued that just as God
could bypass grapes and turn water into wine, so He could bypass
parents. Animals would thus appear directly from ocailta semina, in-
visible seed. Around the year r ooo, Cardinal Pietro Damiani insisted
that birds bloom from fruits, and ducks enlerge from seashells. En-
glish scholar Alexander Neckam (rr 57-rz17) specified that fir trees,
exposed to sea salt, give rise to geese. The Flemish alchemist and

TRUE.SCALE TIMELINE
(Millions of
years ago) 4,600 4,500 4,400
I

I HADEAN EON I

Origin of Earth-Moon Oldest rocks dated by Outgassing of vola-

system and other solar radioactivity from tiles from mantle to
system planets meteor (a chondrite atmosphere
from Canyon Diablo
Abundant impact
crater, Arizona)
cratering
 Once Upon a Planet I gU

physician Jan Baptiste van Helmont (158o-r644) shared his recipe

for making mice from dirry underwear.
-We
moderns may laugh, but the notion of spontaneous genera-
tion made such sense at the time that Gw questioned it. "Since so
little is required to make a being," agreed Descartes, "it is certainly
not surprising that so many animals, worms, and insects form spon-
taneously before our eyes in all putrefying substances."3 Aristotle
had taught that the heat of the male seed animated and formed the
cooler matter carried in the woman's womb. Lacking suflicient male
heat, a woman miscarried or gave birth to a limbless infant. Heat
rnight bypass seed altogether and directly generate worms, bats,
snakes, crickets, or other verrnin from meat or filth- Alchemists used
heat to try to synthesize gold. In a patrilineal, male-dominated Eu-
rope women were like potter's kilns in which the act of fathering
came to ftuition; the female supplied only matter and not the essence
of living form. Even Newton suggested that plants might spring forth
from the coruscation of cometary tails. Nor did the invention of
the microscope sweep away the old idea. Many believed that the
"animalcules"Leeuwenhoek had discovered in plant fluids, ditch wa-
ter, and saliva emerged directly from these fluids, just as veal-left
to its own devices-was thought to generate flies.
Ironically, the notion of spontaneous generation was at first threat-

4,300 4,200 4,'100

I I I

Oldest known mineral Early seas Exuberant volcanism

crystals (present-day
Australia)
Possible existence
of first continents
and meteoric cratering
Oldest rocks dated by
continues
radioactivity of moon
rock
 66 What ls Life?

ened as much by the idea of fixed species as it was by countervail-

ing observations. Species were recorded as fixed categories. The
works of Swedish botanist Carolus Linnaeus (t7o7-r77g), the
founder of modern taxonomy who gave the name Homo sapiens to
the human body (but not to the soul), and those of French anatonrist
Georges Cuvier Q769-r832), who extended the Linnaean classi-
fication to fossils, made the notion of spontaneous generation more
difiicult to accept. For Linnaeus, fixed species were distinct and sep-
arate forms created by an omnipotent God. To Cuvier, fossils are
evidence of past life, in particular of carastrophic floods, at leasr one
of which was recorded in the Bible.
It thus came to be believed that an all-powerful God, once and
for all, had created all Earth's "creatures." Indeed, Swiss naturalist
Charles Bonnet ft7zo-r7y) ruled out spontaneous generation be-
cause it was superfluous to his theory of emboitement-thatthe orig-
inal female of each species had been created, as in a set of Russian
dolls, with the germ cells of all future generations already inside her.
Half-blind Bonnet had discovered the all-female reproduction sys-
tem of certain insects, the parthenogenesis of aphids, a fact which
helped him argue against "svqlusi6n"-the word he used to refer
to the belief of those foolhardy enough to believe in the wanron
notion of species transformation.

4,000 3,900 3,800

I ancrrnu ror* I
I I

Beginning of Earth crust Origins of life in the Creenstone lsua

formation and presumed form of bacterial cells Belt (present-day
start of tectonic activity Creenland), indi-
Appearance of first
Earliest Earth rocks (zircons kingdom: BACTERIA cating possible bio-
from Mount Narryer in (MONERA) logically produced
present-day Australia and carbonate and
Anaerobic prokaryotes reduced carbon
Acasta gneiss from present-
and therefore au-
day northwestern Canada)
topoiesis, metabolism,
dated by radioactivity
and reproduction have
Oldest Mars meteor crater evolved
(estimated)
 once Upon a Planet I e,

"Life from nonlife" was adhered to even after the Florentine physi-
cian and poet Francesco Redi $626-t6g7) performed his diligent
experiments disproving such spontaneous generation. Redi placed
a variety of meats-a snake, some fish, and a slice of veal-in sealed
jars. Another set of jars were left open. Redi's experiment was a
clear success. In his "observations on the generation of insects," he
recorded that he "began to believe that all worms found in meat
were derived from flies, and not putrefaction."4 Redi, in other words,
developed a theory of maggots. Having seen flies hovering around
and entering the open &ut not the closed) jars, he confirmed his
suspicion that the sealed meats, despite their putrid stench, did not
become "wormy." In phase two of the experiment he covered meat
with a cloth that prevented flies from laying eggs. No vermin ap-
peared. He concluded that "Earth, after having brought forth the
first plants and animals by order of the Supreme and Omnipotent
Creator, has never since produced any kind of plants or animals, ei-
ther perGct or imperfect; and everything which we know in past
or present time she had produced, came solely . . . from seeds of the
plants or animals themselves, which thus, through means of their
own, preserve the species."s
Scientists are said to abandon theories as soon as they are contra-
dicted by experiment. In fact, many do the reverse, ignoring awk-

3,700 3,600 3,500

I
I I
I

First appearance of Barberton Mountain Onverwacht Croup and Warra-

banded iron formation Land (present-day South
(BlF), suggesting local Africa) and Pilbara Block
sources of oxygen (present-day Western
at sediment-water Australia), containing
interfaces fossil evidence for an-
oxygenic communities:
microfossils, stromato-
lites, and chemical fossils
woona Croup (present-day South
Africa), containing abundant
reduced carbon in shales, micro-
fossils, and stromatolites, imPlY
widespread occurrence of photo-
synthetic bacterial communities
Earliest direct evidence of tectonic
activity: granite of the Kaapvaal
Craton from present-day South
Africa
 68 What Is Life?

ward experimental evidence in an effort to save appearances. No-

where, to paraphrase Mark Twain, is so much derived from so lit-
tle as in the production of scientific theory from scientific fact. A
century after Redi's experiment, the English naturalist and Roman
Catholic priestJohnTuberville Needham (t7 r 3-r78r) collaborated
with the early evolutionist Georges Louis Leclerc Buffon Glol-
t788). Buffon, as keeper oftheJardin du Roi, the French royal botan-
ical gardens, was author of the forry-four volume ltJatural History,
read by many of the educated class, including Erasmus Darwin (r I I r-
r Soz), Charles's grandfather. Together, Needham and Buffon per-
formed an experiment designed to determine whether spontaneous
generation appLied to all of liG. Boiling mutton broth, they carefully
sealed it in ajar. Opening it a few days later, they saw copious growth,
suggesting to them that spontaneous generation did apply to mi-
crobial life. Although absolutely misleading-because they failed to
kill the boil-proof microbes-the experiment ironically confirmed
Buffon's essentially modern notion that "organic molecules" could
under certain conditions combine to produce rnicroorganisms.
In r768 Italian biologistLazzaro Spallanzant (r729-r799) demon-
strated that his illustrious predecessors Buffon and Needham had
neglected to boil the broth sufiiciently. Still, Spallanzani's tests did
not satisfli Ernst Haeckel, who believed prolonged heat destroyed a

3,400 3,300 3,200

I I I

Development of thickest Trace amounts of Continental tectonic activity-

(and therefore oldest) oxygen gas (O2) many small plates
portlons of continents in atmosphere and
sediments
 Once Upon a Planet 69

"vital principle" in the air. Not until French chemist Louis Pasteur
(t8zz-r895) exposed boiled meat extract to air by means of a flask,
whose long neck was bent down and then up, were vitalists defeated.
Air, but not bacteria, yeasts, or any other sort of life, could rise against
gravity to enter the zigzagpassageway to the life-supporting broth.
As soon as the glass was broken, and microscopic life could enter,
growth on the broth began. No other explanalion held: life came
only from previous life that was begotten by still earlier life. And
yet, the work of Pasteur, proving that life comes only from previ-
ous life, strongly suggested that only God could have created life in
the Beginning.

ORIGINS OF LIFE

In r 87r Darwin mused that one "could conceive in some warm lit-
tle pond, with all sorts of ammonia and phosphoric salts, light, heat,
electriciry etc." a chenrically formed "protein compound . . . ready
to undergo still more complex changes."6To trace liG back to mat-
ter was a logical extension of the idea that all species had evolved
from common ancestor. If species could evolve, what was to stop
a
matter itself from evolving into life?
A youngRussian biochemist, Alexander lvanovich Oparin (r 8S+-

3,100 3,000 2,900

ll I

Formation of Oldest evidence for life in present-day Cold deposited in

Fig Tree Group North America: Steep Rock, Ontario paleoriver in present-
(present-day South day Witwatersrand,
Widespread stromatolite reefs pre-
Africa) of rocks that South Africa, indicating
served at Steep Rock and Pongola Belt
contain microfossils (present-day South Africa) bacterial-mediated
of reproducing cells gold precipitation in
Diversifi cation of bacteria-probably ancient estuaries
all major metabolic modes evolved by
now (e.9., chemoautotrophy such as
Hz, HzS, NH:, and CHa oxidation;
oxygenic photosynthesis; reduction of
iron and manganese oxides to metals)
 70 What ls Life?

r98I), published a book in r929 entitled The Origin of Life Oparin

focused attention on specific r,vays in which chemicals might self-
organize toward life. He described droplets growing by absorbing
carbon compounds in a primeval soup. Theorizing an early hydro-
gen-rich atmosphere with gases such as methane and ammonia, and
a solar source of energy, Oparin postulated that his "coacervates"
or "semiliquid colloidal gels" would become increasingly depen-
dent on their "own specific internal physico-chemical structure."
Eventually,

The internal structure of the droplet determined its abiliry to ab-

sorb with greater or less speed and to incorporate into itself organic
substances dissolved in the surrounding water. This resulted in an
increase in thesize of the droplet, i.e., they acquired the power to
grow. . . . A peculiar selective process had thus corre inro play which
finally resulted in the origin of colloidal systems with a highly de-
veloped physico-chemical organization, namely, the simplest primary
organisms.T

Because Oparin inhabited a nation (the former Soviet LJnion) that

had been ofiicially atheistic since r9r7, he could rheorize on this
new version of spontaneous generation without confronting estab-
lished religion.

2,800 2,700 2,500

I
I I I

Large continents formed Stromatolites abundant and End of major crust-

from raised portion of cosmopolitan on ancient con- forming period
the plates known as the tinents in parts of present-
" pre-Cambrian shields" day Africa, North and South
America, Australia, and Asia
 Once Upon a Planet 71

ln rgzg the British physiologist J.B.S. Haldane published an ar-

ticle making the point that reactive oxygen would have destroyed
organic compounds; developing life, therefore, must have arisen in
an oxygen-free atmosphere.s The work of Haldane and Oparin was
an inspiration to "origins-of-life" experimenters from the lJnited
States, such as Stanley L. Miller, Sidney Fox, and Cyril Ponnampe-
ruma. Nonetheless, Oparin was no more removed from his socio-
cultural milieu than were his predecessors; after World War II he
declared Schrcidinger's book What Is Life? to be "ideologically dan-
gerous" and he protested the new emphasis on genes, viruses, and
nucleic 6alling it "mechanistic." Yet Oparin, by imagining how
".i4t,
life could have first evolved, revived the notion of spontaneous gen-
eration of life from nonlife.
In 1959 organic chemist Sidney Fox and his colleagues cooled
water-free mixtures of amino acids to make "proteinoid micro-
spheres." Resembling cocci bacteria, these microspheres would, un-
der pressure, occasionaUy divide. Leslie Orgel of the Salk Institute
in California discovered a DNA-like molecule (fifty nucleotides
long) that formed spontaneously from simpler carbon compounds
and lead salts. Five years later, ATP-the compound that is univer-
sally used by life to store energy-was produced by Carl Sagan, Ruth
Mariner, and Cyril Ponnamperuma in a phosphorus-containing

2,500 2,400 2,300

I enorenozotc eor I I

Ceologically modern processes Beginning of worldwide Continued

begin: Oxygen gas (Or) begins
to seasonally accumulate; BlFs
conspicuous and abundant;
extensive, huge lakes or oceans;
carbonate platforms, indicating
biogenic reef-like structures made
by bacterial communities in ma-
rine settings
First supercontinent (pre-Pangea)
age of BlFs: 90 percent of expansion of
Earth's current mineable carbonate reef-
iron deposits in present-day like platforms
southern Africa, Brazil, and BlFs
Central America, western
Ontario, northern
Michigan, and Minnesota
formed between 2,400 and
1,8OO million years ago
 72 What ls Life?

mixture of gases thought similar to Earth's early atmosphere. "It is,

perhaps, ironic," writes lJniversity of Maryland chernist Ponnam-
peruma, "that we tell beginning students . . . about Pasteur's ex-
periments as the triumph of reason over ntysticism yet we are con)-
ing back to spontaneous generation, albeit in a more refined and
scientific sense, namely, to chemical evolution."e
The "abiotic production" of ATP was acrually a continuation of
work begun by Stanley L. Miller, a graduate student of Nobel lau-
reate Harold Urey (r893-198r), ar rhe Universiry of Chicago in
t 95 3 . Miniaturizing what he thought was Earth's earliest ambience,
Miller filled flasks with gases (imitation atmosphere) over the sur-
face of sterilized water (imitation ocean). For a week he bombarded
his glassware microcosm with a lightning-li-\e electrical discharge.
The result was the evolutionary biologist's .Diion of the rwitch-
ing limbs of Mary Shelley's Frankensrein monsrer. Alanine and gly-
cine, two chemicals essential to living proteins, as well as many
other compounds, had spontaneously appeared in the flasks. In the
laboratory, cooking from scratch, scientists had thus repeated the pre-
biotic origin not quite of life but of the nutrients needed for self-
maintenance-a sort of primeval food.
Miller's laboratory mock-up of the early planetary atmosphere
contained hydrogen gases like those left over from the gravitational

2,200 2,100 2,000

I t.' I

Widespread lncreasing UV (ultravio- Free Oz abundant in atmosphere, indi-

occurrence of let) ray-absorbing ozone cating dominance of aerobic organisms
prokaryotic shield (Or derived from
Mitochondria, ancestors to most eu-
plankton (bacte- 02 accumulating in
karyotes, acquired by symbiosis as
rioplankton) in atmosphere)
purple eubacteria
world's oceans
Oldest abundant fossil
Cunflint lron Formation (present-day
bacteria: Cunflintia,
Ontario, Canada) and equivalent fossil
Huronospora, Lepto-
biotas in present-day China, Australia,
teichus gol ubici i, etc.
and California containing complex fila-
mentous microfossils and remains of
structured communities
 Once Upon a Planet I ,t

accretion of the sun: hydrogen (H2), water vapor (HzO), ammo-

nia (NH:), and methane (CH+). The experiments showed in star-
tling fashion that the chemicals of life do self-organize without
conscious direction. Given favorable conditions-Miller's model
of the early atmosphere was only a rough guess-organic com-
pounds spontaneously form from simpler precursors. The unde-
niable conclusion was that at least the matter of life spontaneously
generates.
Miller's experiments were repeated and modified by many en-
of energy, such as
thusiastic chernists. Some used alternarive sources
ultraviolet radiation and heat. Akiva Bar-Nun, for example, gener-
ated "sonic booms" in the laboratory; he showed that even energetic
sound waves make protein components from atmospheric gases. Ade-
nine, cytosine, guanine, thyrnine, and uracil-the five nucleic acid
bases that strung together make DNA or RNA molecules-all have
been synthesized in "prebiotic chemistry" experiments.
Of the six kinds of atoms crucial to life on Earth-carbon, ni-
trogen, hydrogen, oxygen, sulfur, and phosphorus-all have been
detected in space. Hydrogen, the most common element in DNA,
RNA, proteins, fats, and other compounds created by life, is also
the most common in the universe. Ammonia (NH:) was discov-
ered in interstellar space in r968. HiCzN, cyanoacetylene, was de-

1,900 1,800 1,700

I I I

First appearance of Replacement of BlFs by Appearance of second

Grypania, identified as red beds (oxidized iron kingdom: PROTOCTISTS
the earliest Protoctista sediments), indicating
Earliest eukaryotes docu-
(maybe reinterpretable worldwide transition to
mented in fossil record as
as discarded cyanobac- an atmosphere rich in
acritarchs, indicating cell
terial sheaths) oxygen
evolution by symbiosis
Origin of speciation inferred
from molecular data on pro-
toctists (primarily anaerobic
mastigote protists)
 74 What ls Life?

tectedin t97o. Alcohol (CH:CHzOH) abounds in the constella-

tion Orion. Other compounds found both in space and in living
things include water, acerylene, formaldehyde, cyanide, methanol
(wood alcohol), and five-atom formic acid, the clear fluid secreted
by agitated ants. The simplest compounds of life form easily by
chemistry alone.
Prebiotic chemist Cyril Ponnamperuma (r9zz-r994) thinks our
planet may have been "knee-deep" in polyaminomaleonitryl, an or-
ganic compound whose combinations could have inaugurated the
later world of cells. Polyaminomaleonitryl is a polymer, a large mol-
ecule made of repeated links of HCN, hydrogen cyanide. HCN, a
simple three-atom compound, has been detected onTitan, Saturn's
sixth (and biggest) moon. A precursor to other biochemicals, in-
cluding adenine and guanine in the nucleotide bases of RNA and
DNA, HCN may be a key ingredient in the cosmic recipe for life.
Ponnamperuma casts it as the "God molecule." The polymer comes
in a range of colors, including the reds and browns characteristic of
"tholins," organic goop formed in the laboratory of astronomer Carl
Sagan at Cornell (Jniversiry under conditions simulating those
thought to exist on the similarly colored clouds of Jupiter.
Meanwhile, British chemist Graham Cairns-Smith has proposed
that clays would have shielded fragile cell precursors from the solar

1,600 1,500 1,400

I I

Diversifi cation of aerobic Protoctist evolution: ori- Appearance of terres-

life gins of mitosis, meiotic trial cyanobacterial
sex, gender, and pro- life (desert crust
Appearance of planktic
grammed death of indi- and soil microbial
and benthic organisms
viduals in eukaryotic communities)
possibly correlated to sym-
microorganisms and their
biotic acquisition of air-
descendants
breathing mitochondria
 once Upon a Planet | ,S

rays which, though capable of assembling organic compounds, could

also irradiate them into oblivion.l0 Crystalline clays, others have sug-
gested, could have aggregated on nonliving bubbles produced by
wind, rain, volcanoes, and waves. Even today, atcracting particles along
their surfaces while undergoing changes in temperature and pres-
sure, bubbles serve as a meeting place where ambient carbon, nitro-
gen, hydrogen, and other elements form more complex compounds.
When bubbles burst they leave chernical residues in their wake.
'Whatever the precise route taken in the origin of life, Freeman
Dyson proposes that it probably came by way of a kind of molec-
ular "symbiosis" (although the word is not quite right, since nei-
ther partner was itself alive) between RNA-a "supermolecule"
probably crucial, as we shall see, to life's origin-and more hap-
hazardly growing "protein creatures"ll Despite much conjecture
and intriguing research, it must be remembered that no life has yet
been synthesized in the laboratory. The gap berween chemical evo-
lution (the appearance of carbon compounds by lipids or fatry films
that appear in "environmental" mixtures) and true cells (self-
bounded, self-maintaining, and ultimately reproducing matter) re-
mains unbridged. Nonetheless, the laboratory-explored shenani-
gans of RNA-as we shall see in a moment-are shrinking this gap
daily.

1,300 1,200 1 ,100

I I

Diversification of sea- Continued diversifi cation Clobal rifting event

weeds (algae, which are and widespread appear-
photosynthetic protoc- ance of monera (iron
tists) of unknown taxa bacteria, cyanobacteria,
possibly correlated to and many unknown
symbiotic acquisition of Protoctista-sexual cysts,
photosynthetic plastids algae, microscopic and
even large fossils)
 75 What ls Life?

"STUMBLING FORWARD"

Our scientific understanding today of the origins of life is proba-

bly no better than was our understanding fiffy thousand years ago
of the origins of fire. 'We can maintain and play with it, but we can't
yet start it. The assumption that the origin of life may be repeated
by investigators in the laboratory is a shocking example of the au-
daciry of scientists-and yet it may prove correct.
Scientific investigation does reveal gradations between certain
chemical systems and the animated material all of us recognize as
life. Schrcidinger's crystal analogy has given way to an idea of life as
a chemical system requiring material and energy to persist far from
thermodynamic equilibrium, i.e., a dissipative system. Dissipative
systems that are not alive may nevertheless act in ways that are eerily
lifelike. One such dissipative system develops in the Belousov-
Zhabotinsky reaction. It involves the oxidation of malonic acid by
bromate in a sulfuric acid solution containing cerium, iron, or man-
ganese atoms (see plates 5a and 5b). Under certain conditions, con-
centric and rotating spiral waves will occur in an aesthetically cap-
tivating chemical reaction that may last for hours.
The regularity and duration of such reactions have led some sci-
entists to compare them with life. Using energy from outside to in-

1,000 900 800

I I I
lncrease in diversity of Oldest " giant" acantho- Worldwide proliferation
algae (photosynthetic morph acritarchs, prob- of unidentified large
eukaryotes) and other ably algae "quilted " organisms,
protists fossilized in sandstone,
probably colonial sand-
dwelling members of
diverse protoctist king-
dom, the Ediacaran biota
 Once Upon a Planet 77

of which
crease their internal order, these chemical systenN, some
arebrightly colored, "live" for a while beyond the limit of equilib-
rium chemistry. ErichJantsch (r9zo-r98 j), an Austrian-American
astrophysicist and philosopher, explains that,

Whereas free energy and new reaction participants are imported, en-
tropy and reaction end products are exported-we find here the rne-
'With
tabolism of a system in its simplest manifestation. the help of this
energy and matter exchange with the environment, the system main-
tains its inner non-equilibrium, and the non-equilibrium, in turn,
maintains the exchange processes. One may think of the image of a
person who stunrbles, loses his equilibrium and can only avoid falling
on his nose by continuing to stumble forward. A dissipative structure
continuously renews itself and maintains a particular dynamic r6gime,
a globally stable space-time structure. It seems to be interested solely
in its own integriry and self-renewal.12

Dissipative structures, chemical systems that use streams of energy

to increase their internal order, are, however, rare and short-lived.
But if the increased internal order is that of life, then, given access
of the system to a source of energy and the right kind of matter
(nutrients), it maintains indefinitely. This is autopoiesis. Autopoiesis
is what happens when a self-bounded chemical system-based not
on small molecules of sulfuric and malonic acids but on long-

(Beginning of "Human-centered" Timeline)

700 600 541

I I I exnmnozore eoru

Seriesof worldwide Appearance of third Paleozoic Era

"Vendian" ice ages kingdom: ANIMALS
Cambrian. Ordovician. Silurian.
followed by diverse lnferred origins of egg, and other time-rock divisions
new planktic and (see page 53)
sperm, embryo, and
benthic communi-
blastula
ties, probably of "Age of Marine Animals"
protoctists Appearance in fossil
Trilobites and other hard-shelled
record of soft-bodied
animals prevalent
animals (sponges, coe-
lenterates, arthropods, Appearance of fourth and fifth
and others) kingdoms: PLANTS and FUNGI
 78 What ls Life?

molecule nucleic acids and proteins-reaches a critical point and

never stops metabolizing.
The cell, the smallest autopoietic structure known today, is the
minimal unit that is capable of incessant self-organizing metabolism.
The origin of the tiniest bacterial cell, the first autopoietic system
with about five hundred different kinds of proteins and other long
chain molecules, is obscure. Yet most agree that complex carbon
compounds, exposed in some way to unceasing energy and envi-
ronmental transformation, became oily droplets that eventually be-
came membrane-bounded cells.
Metabolism, the chemical measure, the specific earttrly manifes-
tation of autopoiesis, has been a property of life since it began. The
first cells metabolized: they used energy (from light or from a small
range of chemicals-never from heat or mechanical movement) and
material (water and salts, carbon, nitrogen, and sulfur compounds)
from outside to make, maintain, and remake themselves. Auto-
poiesis, the chemical basis for the impatience of living beings, is never
optional. Absolutely required at all times for any life form in a wa-
tery milieu, autopoiesis, once it appeared in the tiniest bacterial an-
cestor, was never completely lost.
You embody the processes of the early Earth in your living cells.
The failure of the autopoietic system of cell maintenance is death.

500 400 300

t' t
 Once Upon a Planet I ,,

If autopoiesis of a cell ceases, the cell dies. A many-celled organ-

ism capable of replacing its cells survives as the autopoietic behav-
ior of the larger organic being prevails. If too many component cells
die, metabolism of the larger entiry halts and death follows. Any cell
or organism that continues to self-maintain will grow, and the im-
perative to reproduce will follow. Although not obvious to the naked
eye, cell metabolism never stops. Chemical transformations such as

nutrient uptake and energy conversion, and the fabrication of

DNA, RNA, and proteins, occur continuously in all cells and all be-
ings made of cells.
Life seems to have originated in whatever were the primordial
ancestors of modern bacteria. Chemical systems that became bio-
logical systems, these first beings would have metabolized and in-
corporated energy, nutrients, water, and salts into their developing
selves. The first cells formed. As in Jantsch's analogy of the person
stumbling forward to avoid falling on his face, so membrane-
bounded cells replicaringRNA and producing other molecules stum-
bled onto DNA-based RNA and protein synthesis; that is, repro-
duction became a means to retain self-maintenance, to postpone a
return to thermodynamic equilibrium.
Bacteria reproduce in the time it takes to read this chapter. Ele-
phant and whale reproduction can require a decade. But whatever

245 200 100

I rnesozorc rna I I

Triassic, Jurassic, and

Cretaceous periods
"Age of Reptiles"
 80 What ls Life?

the rate, reproduction requires DNA replication in cells. It requires

RNA, protein, and membrane synthesis and the intrinsic locomo-
tion of growth. Reproduction of larger beings-protoctists, fungi,
animals, and plants-also involves growth and division of their com-
ponent cells. Autopoietic multicellular beings are composed of cells
which themselves are autopoietic. Animal and plant reproduction
is a permutation of cell autopoiesis, just as cell autopoiesis is a per-
mutation of nucleic acid and protein metabolism. Our instinctive
desire to live is directly related to the autopoietic imperative to sur-
vive, itself related to the "yearning" of heat to dissipate.

METABOLIC WINDOWS

Because ceUs retain their organization in spite of-or because oF-

the helter skelter around them, they provide science with a window
onto the past. It is a rather magical fact that, within the autopoietic,
thermodynamic view, our bodies today should have virtually the
same chemistry as that prevailing on Earth's surface 3,ooo rnillion
years ago. Remember that when life became autopoietic it postponed
indefinitely the moment of total heat equalization and loss of or-
der. Using the energy of food and sunlight, life has thwarted ther-
modynarnic equilibrium.

654
*..,..,
I ceruozotc-e*l l eresent
"Age of Mammals"
|iffiil::*
ancestors
 Once Upon a Planet I t,

Death is illusory in quite a real sense. As sheer persistence of bio-

chemistry, "we" have never died d.uring the passage of 3,ooo mil-
lion years. Mountains and seas and even supercontinents have come
and gone, but we have persisted.
We have, of course, had to "up the stakes" at various junctures
to stay alive. This continuous "upping of the stakes," which, on the
personal level, links desire to death, is on the species level described
asevolution. Beings always require food and energy to stay the same,
and often they have to evolve, to change into new forms, simply to
self-maintain. The feline lineage, the flowering plant lineage, the
nautiloid-squid and the rest of the cephalopod lineage have changed
and persisted through the sexual reproduction and death of their
members.
Evolution, no less than the nucleic acid replicarion of autopoiesis
and reproduction, is a "stumbling forward" to stave offthe threat of
thermodynamrc dissolution. Most atoms in our bodies are hydrogen-
the element which as a gas was, according to astronomic models,
blasted beyond the confines of the inner solar system when the sun
turned on. Nonetheless, these atoms which should be long gone
have defied time and space by becoming bound up with (a$ life.
Today hydrogen-rich gases such as amrnonia exist not only in the
atmospheres of the giant outer planets but in the inner solar system
where life has preserved them in its self-sirnilar structure ever since
it began maintaining and reproducing.
Indeed, the original dissipative chemistry the protein and nucleic
acid chemical clocks that arose prior to life may even have been pre-
served. One of the most beautiful aspects of living things is that they
bear within their very form the presence of the past. We resemble
our parents and other people who lived ten thousand years ago. This
preservation of the past in the present is fortunate for scientists. Each
body is the charitable gift of a biochemical museum, and each bac-
terial cell an unplanned time capsule.
Far from lost in what Shakespeare called the "dark backward and
abysm of time," liG's origins are an open secret awaiting decipher-
 What ls Life?

ing by sufirciently talented chemists. If life is an autopoietic, far-

from-equilibrium phenomenon, living cells should srill contain
significant fragments of preliving systems. Vestiges of life's origin
may still exist, a stuttering genesis for scientists patient enough to
listen. Life may even contain the original dissipative srructures and
chernical fossils in the form of metabolic pathways. Ultimately far
more valuable than microfossils, or the modern alchemical experi-
ments of energizing chemicals in laboratory glassware, are organ-
isms: easy to overlook in their obviousness-uncannily present-
they are metabolic windows onto life's origin.

THE RNA SUPERMOLECULE

The minimal free-living autopoietic endry roday is probably a tiny,

spherical, o>'rygen-shunning bacterium which requires energy and
food to keep going its five hundred genes and proteins. Or maybe
it is a kind of mycoplasm, a being so small that until recendy it was
known only as a cousin to a growing speck that caused disease symp-
toms in brains of sheep. Even in these, the bonded atoms of car-
bon, hydrogen, nitrogen, and oxygen interact recursively in a meta-
bolic system.
The genes that are DNA require active RNA to work. DNA and
RNA together make the proteins that form cell structures, and they
also make the very enzymes that slice and splice the genes. The so-
called genetic code actually refers to the correspondence berween
the linear order of DNA's components and that of the amino acids
in a vast array of different proteins.
With the help of RNA, the nucleotides of DNA line up amino
acids of protein. Our blood, internal organs, fingernails, skin, and
hair are all made of proteins. The reason that nutritionists advise us
to eat the "eight essential amino acids" is because the human body
cannot renew itself without taking in these protein components from
food and drink. The human body cannot synthesize these particu-
Iar amino acids at all-even from their simpler components.
 once Upon a Planet ] ta

In contrast to our human need to mine the environment for es-

sential amino acids, no living being on Earth needs to stalk the en-
vironment in quest of the deoxyribose sugar essential for its DNA.
Rather, deoxyribose is synthesized in cells by diverting an oxygen
atom from ribose. It is ribose, the five-carbon sugar of RNA, that is
often taken up from the outside as food. That all cells, given ribose
sugar, can make deoxyribose from it suggests that ribose came first.
RNA, with ribose, evolved before DNA. DNA sugar metabolism
evolved by subtraction of oxygen from RNA sugars. The earliest cells
may have been RNA beings which only later evolved DNA systems.
Comparing RNA and DNA metabolism is an example of peering
into cellular windows for clues to life's most ancient origins.
Other evidence questions the pretensions of DNA, the "master
molecule" to life's biochemical throne. RNA, more versatile than
DNA, is a better choice for the replicative tool of life's earliest au-
'Whereas
topoietic system. double-stranded DNA uses deoxyribose
sugar for its chain, single-stranded RNA uses ribose, the source ma-
terial of deoxyribose. Unlike DNA, which must use RNA to code
for proteins, RNA by itself can direct both its own replication and
the making of proteins. In ancient times, RNA probably did all that
DNA does today inside of cells, and more. In all cells, when the
rwo helically coiled strands of DNA open to expose a section of the
nucleotide sequence, that portion of the DNA is "copied" onto
nressenger RNA. Taking its message to r\ivo other kinds of RNA
(transfer RNA and ribosomal RNA-named for the ribosomes, the
"factories" in the cell where proteins are made), messenger RNA's
information is "translated" into the amino acid units that assemble
into working proteins. RNA can make proteins, in principle, with-
out any DNA.
Followrng the lead of Sol Spiegelman (r9r4-r983) at the lJniver-
sity of Illinois in the late r96os, German Nobel physicist Manfred
Eigen (together with coworkers at the Gottingen Institute) found a
way to induce test tube RNA molecules to replicate by themselves.
Eigen showed that nucleotide units of RNA lined up and formed
 84 What ls Life?

functional RNA. Most impressively, some of the test-tube RNA even

mutated into a different RNA that replicated more quickly than the
original. The Eigen experiment did not, of course, reveal the spon-
taneous generation of life; RNA molecules by themselves are not
cells. RNA in test tubes would have remained completely lifeles had
scientists not extracted proteins from live cells and added them to
test tubes containing RNA.
Eigen's RNA molecules are much like viruses. Certainly not
alive, they show a power on the border of life. As computer viruses
require human-run computers to spread, so naturally occurring
viruses-not full, autopoietic beings but genes coated with protein-
require living cells. Replicating RNA viruses may be just as dan-
gerous and capable of replication as are DNA viruses.
Donald Mills of Columbia Universiry also made test-tube RNA
viruses; these RNA viruses used a bacterial enzyme to replicate
themselves inside a bacterium Mills had conveniently provided for
them. In the early r98os Thornas Cech of the lJniversiry of Col-
orado and Sidney Altman ofYale Universiry found that certain kinds
of RNA are capable of self-splicing.. In recognition that RNA acts
like its own active protein by cutting and rearranging itself the way
enzyme proteins do, these test-tube reactants are called "ribozyme5."
Ribozymes, cell-free nrixtures of the right pieces of RNA, proteins,
and their components, furthermore change with time in a kind of
test-tube evolution. Gerald Joyce at the lJniversiry of California,
San Diego, may be on to the hottest biochemical project yet: with
his colleagues Jack'W. Szostak and A. D. Ellington of the lJniver-
sity of Indiana, Bloomington, he has found ribozymes for which
the associated enzymes actually accelerate the rate of RNA replica-
tion: veritable molecular evolution in a test tube.13
RNA is thus a prime candidate for early life's supermolecule. By
working on itself as it grew, RNA may have generated a m6lange
of expanding possibilities. Able to replicate and mutare, acting as
both enzyme and gene, RNA performs operations that form more
of itself.
'We
can now envision that on the Archean Earth it was bounded
 Once Upon a Planet 85

RNA that developed replication and information systems, as. se-

questered inside oil droplets, it began the trek toward autopoiesis.
The DNA world of all life today may thus have evolved inside RNA
cells of the "RNA world," a term coined in I986 by Harvard Uni-
versiry biologist and Nobel laureate Walter Gilbert.la

CELLS FIRST

In the normal waking state, human bodies burn sugars aerobically,

using oxygen atoms drawn from the air. But in strenuous activify
the body reverts to a distinct metabolism; muscles ferment sugars
in the same anaerobic way invented by early bacteria. When stressed,
our bodies thus "remember" the times before the atmosphere became
suffused with oxygen. Such physiological flashbacks re-present past
environmental conditions and the bodies that evolved to live in them.
In a very real sense, all beings today retain traces of Earth's earliest
biosphere.
Neither DNA nor RNA alone is enough to form life. American
biophysicist Harold Morowitz, using the concept that "metabolism
recapitulates biogenesis" ("biogenesis" referring to the origin of life),
suggests that nearly all aspects and forms of metabolism and the syn-
thesis of proteins and nucleic acids evolved only after membranes
enclosed the precursors of cells. Whether proteins or nucleic acids
"came first" in the sequence of life's origins, membranes surely arose
even earlier. Life is thus truly a cellular phenomenon.
Morowitz emphasizes that, in the watery environment in which
life arose, a nonaqueous barrier was necessary to separate the cell
from its surroundings. "To be an entiry distinguished from the en-
vironment, requires a barrier to free diffusion. The necessity of ther-
modynamically isolating a subsystem is an irreducible condition of
Iife. . . . It is the closure of an amphiphilic bilayer membrane into a
vesicle that represents discrete transition from nonlife to life."1s Con-
sidered materially, as a system of matter and energy, life is recog-
nizable by its partial separation from the environment by way of a
membrane.
 86 What ls Life?

Over evolutionary time, individualiry always based on the mem-

brane-bounded unit of the cell, arises at ever greater levels of in-
tegration. Life not only evolves, it epitomizes evolutionary transi-
tions. A cell membrane is in common and necessarily intact in all
autopoietic beings. A membrane is a precondition for cell metab-
olism. The first autopoietic system, which may have lacked both
DNA and RNA, was almost certainly a cell. It may have become an
RNA-filled cell, a lipid capsule whose autopoietic persisrence. en-
hanced by the addition of DNA, evolved into our earliest bacterial
ancestor.
At the beginning of this chapter we wrote that life had been aban-
doned by the world, yet the world had gone nowhere. We hope to
have shown that this was more than poetic metaphor: life was in the
midst of the world and matter but separated from it by a translu-
cent, semipermeable membrane.

SO, WHAT lS LIFE? Life is the representation, the "presencing" of

past chemistries, a past environment of the early Earth that, because
of liG, remains on the modern Earth. It is the watery, membrane-
bounded encapsulation of space-time. Death is part of life because
even dying matter, once it reproduces, rescues complex chemical
systems and budding dissipative structures from thermodynarnic
equilibrium. Life is a nexus of increasing sensitiviry and complex-
iry in a universe of parenr matter that seems stupid and unfeeling
in comparison. Life must maintain itself against the universal ten-
dency of heat to dissipate with time. This thermodynamic view ex-
plains, in a way, the determination, the purposefulness of life-for
billions of years it has been stuck in a pattern wluch, even if it wanted
to, it can't get out of, of upping the stakes as it goes. For life itself
is these patterns of chemical conservation in a universe tending to-
ward heat loss and disintegration. Preserving the past, making a
difference berween past and present, life binds time, expanding com-
plexiry and creating new problems for itself.
 MASTERS OF THE BIOSPHERE

Had [bacteria] been discovered on Mars, their description would

have been much more dramatic and the bizarre quality of their
natural history, which often seems like science fiction, would not
have been missed.

SORIN SONEA AND MAURICE PANISSET

Perhaps some of my readers will respond with a smile to my

doctrine of living contagions.
AGOSTINO BASSI

FEAR OF A BACTERIAL PLANET

Microorganisms were a curiosiry a kind of natural history sideshow,

before it was realized that some cause disease. Antoni van Leeuwen-
hoek, inventor of an early version of the microscope in the r67os,
described these beings as "animalcules"-finy animals. He was struck
by their rapid movement, odd shapes, and sheer quantity. In r83r
the half-blind Italian law student Agostino Bassi (r773-r856) proved
the existence of infection by spreading muscardine (silkworm dis-
ease) from one fungus-infected worm to another. Nonetheless, a
generation after Bassi showed that disease did not arise spontaneously,
even Pasteur thought of bacteria only as agents of decay.
A turning point occurred when Robert Koch (r 843-r 9 r o) found
bacteria in blood of cows stricken by anthrax. These little rods
("bacilli") grew from hardy bacterial spores. Feeding them blood
serum, Koch, a German medical ofiicer, learned to grow the bac-
teria in a liquid broth. He developed a stain for them, photograph-

87
 88 What ls Life?

ing the culprits by mounting a cameraon a microscope. Yet the now-

cofilrnon notion that bacteria caused infectious disease was slow to
be accepted. English nurse and philanthropist Florence Nightingale
(r8zo-r9ro) denied the existence of germs to her death.
When the germ theory of contagion finally caught on, it did so
with a vengeance. Different types of bacteria were implicated in an-
thrax, gonorrhea, ryphoid, and leprosy. Microbes, once amusing lit-
tle anomalies, became demonized. Pasteur, like Howard Hughes af-
ter him, had a phobia about dirt and germs. He avoided handshakes.
Wiping down his crockery, he meticulously sought evidence of
wood, wool, and other detritus in his food. No longer an amusing
scandal for parlor conversation, microbes became a virulent "other"
to be destroyed. The metaphor of tenacious, infectious bacteria was
used in Nazi rhetoric of genocide. Today the low regard for bacte-
ria as Lilliputian "agents of disease" still obscures their enormous
importance to the well-being of all the rest of life.
Until the rg jos the oldest unambiguous fossils were 5zo-million-
year-old trilobites and other extinct marine animals. The oldest rocks
on Earth, by contrast, dated to nearly 4,ooo million years. Fossils of
microorganisms have now been confirmed in the oldest sedimen-
tary rocks, suggesting that life took root soon after the Earth-Moon
system formed.
ln 1977 paleobiologists Elso Barghoorn and Andrew Knoll of
Harvard University found some two hundred fossil bacteria (some
even in stages of cell division) in sedimentary rock 3,4oo million
years old. Because Barghoorn had earlier discovered microbial life
in the Gunflint Iron Formation of western Ontario and states bor-
dering Lake Superior, he was well prepared to detect the fossil re-
mains of bacteria in these vastly older African and Australian rocks.
In r99o geologist Maud Walsh went ro the Barberton Mountain
Land in southern Africa and colleced ancient black rocks called
cherts, silica-rich rocks petrified from mud flats and volcanic pools.
Back in the Baton Rouge, Louisiana, laboratory, the cherts were
sliced and polished into thin sections for examination under a mi-
 Masters of the Biosphere I t,

croscope. Walsh saw more than just bacteria; she saw bacteria
trapped in sandy layers of their own making, testifying to whole
microbial mat corrununities that had flourished 3,5oo million years
ago.l
Modern bacteria may offer even more revealing clues about ear-
liest life. American molecular biologist Carl Woese has found that
three rypes of very tough bacteria are distinguished from all others
by their ribosomal RNA: salt-loving "halophiles," heat-loving "ther-
mophiles" of hot springs, and methane*producing "methanogens."
These extreme-condition dwellers have ribosomal DNA that makes
them more similar to one another than to all other bacteria. 'Woese
calls the hardy beings "archaebacteria"; he suggests they are direct
descendants of earliest life on Earth.
The observation that archaebacteria inhabit oxygen-free envi-
ronments-such as the ocean bottom, stomachs of cows, oxygen-
depleted sewer water, and the hot acidic springs of Yellowstone
National Park-agrees with the contemporary picture of a hot
Archean Earth with, at most, traces of oxygen in the atmosphere.
Oxygen was only released into the atmosphere once blue-green
bacteria evolved a way to use energy from sunlight to break apart
water molecules (HzO) to grab their precious hydrogen. Com-
bining the hydrogen with carbon atoms drawn from then-abundant
carbon dioxide, blue-green bacteria were able to manufacture
DNA, proteins, sugars, and all their other cell components. These
light-needy bacteria quickly expanded to sunny waters every-
where on the Archean Earth. In so doing, they released vast
amounts of molecular oxygen left over from their hydrogen min-
ing of water.
Earth's atmosphere thus became an extension of the metabolism
of evolving bacteria. Only through the workings of the most in-
novative bacteria of all time did the originally anoxic Earth gain an
oxygen-rich atmosphere. The planet had first been populated by
methane-makers, sulfur-lovers, and other anaerobes-beings that
neither produced nor used oxygen gas in their rnetabolism.
 90 What ls Life?

LIFE IS BACTERIA

One legitimate answer to the question "What is life?" is "bacteria."

Any organism,if not itself a live bacterium, is then a descendant-
one way or another-of a bacterium or, more likely, mergers of
several kinds of bacteria. Bacteria initially populated the planet and
have never relinquished their hold.
Bacteria may be Earth's tiniest life forms, but they took giant steps
in evolution. Bacteria even invented multicellularity. Most bacteria
in nature, in spite of popular belief, are neulticellular (see plate 6).
In these multicellular bacterial beings, each unit cell itself is a bac-
terial cell.
Some lineages of bacteria went on to evolve into many different
kinds of beings, including ourselves. Inside the cells of all of us right
now are former bacteria, using oxygen to generate energy. These
are the rnitochondria. The photosynthetic blue-green beings and
their descendants (plastids of plants) remove carbon dioxide from
the atmosphere, using carbon for their bodies and eliminating as
waste the oxygen of fresh air-only a small portion of which is used
by the mitochondria that cohabit with what were once photosyn-
thetic bacteria in all plant cells.
Our neighbor planets, Mars and Venus, both have atmospheres
more than 90 percent carbon dioxide. Yet Earth's air is a reactive
mixture, with its huge portion of oxygen and less than o.r percent
carbon dioxide. It was bacteria that removed the carbon dioxide and
produced the oxygen. Bacteria, truly, have made the planetary en-
vironment what it is today. Al1 larger beings contain mitochondria
within their cells, living descendants of bacteria that lived on Earth
before oxygen had accumulated in the air. Life on Earth is a hol-
archy, a nested fractal network of interdependent beings.
The fear of bacteria is, in a way, a fear of life, of ourselves at an
earlier stage of evolution. It is not surprising that microbes now find
us so attractive. Because the carbon-hydrogen compounds of all or-
ganisms are already in an ordered state, the human body (like that
 Masters of the Biosphere 91

of every other living being) is a desirable food source for these tiny
life forms. Bacteria seek us as a source of autopoietic maintenance
in their ancient struggle against thermodynamic equilibrium.
Perhaps we should take solace in the fact that the matter of our
bodies returns upon death not to an inert state of matter but to the
bacterial order undergirding the biosphere. "Don't you see," wrote
Giordano Bruno, "that which was seed will get green herb, and herb
will turn into ear and ear into bread. Bread will turn into nutrient
liquid, which produces blood, from blood semen, embryo, men,
corpse, Earth, rock, and mineral and thus matter will change its form
ever and ever and is capable of taking any natural form."2
The wish to maintain one's youth, one's most attractive form, and
ultimately one's very life is thwarted at the level of the animal body.
But our individual defeat is a victory for the bacteria, which return
the hydrogen-carbon compounds of our bodies to a living envi-
ronment. Closer to life's original structures, bacteria do not live as
we live, toward death. Banning an unfortunate accident, a muta-
tion, or a gene-trading encounter with another bacterium, a single
bacterial cell can "survive" essentially forever in its original form,
as generation after generation of bacterial copies of itseH are made
by cell division.
We pluricellular creatures are each a disequilibrium structure of
cells, as a bacterium is a disequilibrium structure of matter. Hu-
manity as a species, even the entire kingdom of animals, has a far
more fragile existence than theirs-just as their existence is more
tenuous than that of nonliving matter.

THE METABOLICALLY GIFTED

Bacteria can swim like animals, photosynthesize like plants, and cause
decay like fungi. One or another of these microbial geniuses can
detect light, produce alcohol, waft hydrogen and fix nitrogen gas,
ferment sugar to vinegar, convert sulfate ions or sulfur globules in
salt water to hydrogen sulfide gas. They do all this and much more
 92 What ls Life?

not because they are "pathogens" or in service to clean our human

environment but because their survival imperative led to their in-
venting every major kind of metabolic transformation on the
planet.
The smallest of them have a diameter only a thousand times
greater than a hydrogen atom. If there were such a thing as angels
that could dance on the head of a pin, bacteria would be they. An-
cient bacteria mastered nanotechnology. Already miniaturized,
bacteria control specific molecules in ways of which human engi-
neers can only dream. Far more complex than any computer or
robot, the common bacterium perceives and swims toward its food.
Choosing and approaching their destinations, bacteria propel them-
selves by flagella, corkscrew-shaped spinning protein filaments at-
tached to living motors in the membranes of their cells. Conplete
with rings, tiny bearings, and rotors, they are called "proton mo-
tors" and spin at about r5,ooo rpnt. These proton motors move
bacteria in the same way that "electric fan" outboard motors propel
boats.
Rapidly reproducing, bacteria properly supplied with food and
water double their cells in a half hour or faster. They have been
and probably always will be the most importanr players in rnain-
taining the biosphere. A single photosynthetic blue-green bac-
terium growing and dividing under ideal conditions could, in the-
ory, produce all the oxygen now in the atmosphere in just a fer.,r,
months.
All other life forms depend on the workings of uncountable liv-
ing, dying, and metabolizing bacteria. Our relations to the bacte-
ria all around us factor into our health and well-being and rhat of
our soil, food, and pets. As naked and simple as bacterial cells seem,
they are continuously busy at the cellular and the planetary level.
Varieties of bacteria exist that photosynthesize without ever re-
leasing oxygen, yet they use carbon dioxide and hydrogen to pro-
duce all their cell parts. Others incorporate carbon dioxide and hy-
drogen from the air into body protein, converting their waste into
methane gas. Still others turn sulfate to sulfide, or incorporate in-
 Masters of the Biosphere I gS

ert nitrogen into their bodies. Only citizens of the bacterial king-
dom are so metabolically gifted. 'W'hen an anineal (like the terndte
who produces methane) or a plant (like the starved bean who be-
gins to supply itself with nitrogen from its roots) is discovered with
such metabolic skills, it is because they have co-opted the bacterial
bodies to their expertise. Such borrowing also applies to biotech-
nology performed by humans in white lab coats.'We hurnans do
not "invent" patentable microbes through genetic recombination;
rather, we have learned to exploit and manipulate bacteria's ancient
propensiry to trade genes.

THE GENE TRADERS

Bacteria trade genes more frantically than pit full of commodities

a

traders on the floor of the Chicago Mercantile Exchange. The trad-

ing by bacteria of genetic information provides the basis for un-
derstanding new concepts of evolution.
Evolution is no linear family tree but change in the single, mul-
tidimensional being that has grown now to cover the entire surface
of Earth. This planet-sized being, sensitive from the beginning, has
become more expansive and self-reflexive as, for the past 3,ooo mil-
lion years, it has evolved away from thermodynamic equilibrium.
Imagine that in a cofGe house you brush up against a grry with green
hair. In so doing, you acquire that part of his genetic endowment,
along with perhaps a few more novel items. Not only can you now
transrnit the gene for green hair to your children, but you yourself
leave the coffee shop with green hair. Bacteria indulge in this sort
of casual, quick gene acquisition all the time. Bathing, they release
their genes into the surrounding liquid. If the standard definition
of species, a group of organisms that interbreed only among them-
selves, is applied to bacteria, t o

%ftnf;lfrry. The Archean Earth was a promiscuous place of

prodigious growth and rapid gene transfer that led, by and by, to
the genetic restrictions of the Proterozoic protists, the larger com-
posite beings presented in chapter 5.
 94 What ls Life?

Unlike all familiar sexually reproducing species, whose members

have cells with nuclei in them that package their DNA, the DNA
of bacteria is loose inside their bodies. Bacterial cells entirely lack
nuclei; for this reason bacteria are prokaryotes composed of prokary-
otic cells. "Prokaryote" literally means "before nuclei." Free of nu-
clei, and unfettered even by the red-staining, protein-coated chro-
mosomes of all other forms of life, bacteria never reproduce by
mitosis. Mitosis, the "dance of the chromosomes," is the kind of
cell division by which the cells of plants, fungi, and animals always
divide. This dance evolved in protists of the Proterozoic, the eon
that followed the Archean. By contrast, a parent bacteriunt elon-
gates its DNA, dragged by growing membrane to which it is at-
tached, until the full-grown cell splits to form nvo offspring iden-
tical to it. Some bacteria reproduce by "buds," protrusions on the
single parent that yield smaller offspring, all of which contain the
parent's same genes.
Members of familiar species of plants and animals reproduce "ver-
tically," mother and father each donate an equal number of genes
as

(on chromosomes) to form new offspring. Bacteria are under no

such constraint. Rather, bacteria trade genes "horizontally," acquir-
ing new genes from peers in their own generation.
Bacterial cel1s often have spare strands of DNA, that is, extra sets
of genes. These genes may be traded in naked pieces called plasmids
or as protein-coated pieces called viruses. In some bacteria a cell
bridge forms between the one donating its genes and the one re-
ceiving them (fig. 6). This process of growing a cell bridge through
which genes are sent, called conjugation, is distinct from mammalian
sex. No bacterial cells fuse nor do "parents" make equal contribu-
tions to an offspring. Rather, one bacterium, the "donor," passes its
genes in one direction to the "recipient," which does not recipro-
cate the favor. Still, this conjugation meets the minineal requirements
of a biological sex act since the transfer of genes prqfluces a new
bacterium, a "genetic recombinant" being with genes from more
than a single parent.
 Masters of the Biosphere lru

FIcURE 6. Three-way genetic exchange among bacteria. Unlike all otherforms of

life on Earth, bacteria transmit genetic information relatively freely, such that taxo-
nomically different "species" can trade genes. Bacterial sex, important to the evolu-
tion of cells with nuclei (eukaryotes), was probably rampant before bacteria them-
selves produced sufficient oxygen gas to create an ozone layer. The male, on the right
side of the electron micrograph, sends genes through two tubes covered by bacte-
riophage viruses. Kingdom: Bacteria (Monera). Phylum: Proteobacteria.

Bridge-forming bacterial conjugation is limited. Many rypes of

bacteria that cannot conjugate indulge in viral or plasmid sex. Those
that practice this more common form of sex require a difference
berlveen the bacterial "genders": a donor needs a recipient.'Whether
any given bacterial cell is donor or recipient is determined by a sin-
gle "sex" gene. The sex gene may itself be transferred in the con-
 What ls Life?

jugation process. If this occurs, a "male" (donor) bacterium can be-

come a "female" (recipient). "She" becomes a male donor Like "him-
self." Any number from a very few to a very many genes may be
transferred at a time, conferring on the recipient not just an ability
to make cell bridges but other useful traits, such as an abiliry to man-
ufacture vitamins or to resist a particular antibiotic.
When exposed to ultraviolet radiation, healthy bacteria explode
with tiny viruses called bacteriophage. Such viruses spread genes to
surviving recipients. Because on the early Earth atmospheric ozone
was not around to intercept the sun's ultraviolet rays, genetic ex-
change may have been even more prevalent then than it is today.
The early UV-bombarded Earth may have been the scene of a
multimillion-year orgy of gene-trading bacterial sex.
Bacterial recombination is a natural form of the "genetic" re-
combination exploited by biotechnologists. Manipulating a preex-
isting bacterial penchant, technicians force the colon bacterium Es-
cherischia coli to produce, for example, human insulin. The bacterium
takes up a particular human gene, reproduces, and the population
then makes large quantities of this human hormone, normally pro-
duced by the pancreas.
DNA may separate from one dying bacterium and, either as pure
DNA or coated with protein in a viral particle, it may splice into
the genes of another bacterium. Unlike egg and spernl, bacterial
cells never fuse. Only their genes flow, but this flow connects them
into a living planetary genetic miasma, which would probably win
a Hugo Award were it presented as science fiction. Imagine you
are a blue-eyed person (perhaps with newly acquired green hair)
who, in a swimming pool, gulps the more conlmon gene for
brown eyes. Toweling ofl you pick up genes from sunflowers and
pigeons. Soon the brown-eyed you is sprouting petals and flying-
eventually reproducing into gliding brown-eyed, green-haired quin-
tuplets. This fantasy is mundane reality in the world of bacteria,
except that most genes traded there are for metabolic and subvis-
ible traits.
 Masters of the Biosphere I g,

OUR SPLENDID KIN

More species of beetle inhabit Earth than any other kind of life, but
bacteria are by far the most numerous organisms on Earth. Thken
together, bacteria are also the most diverse. They are the oldest, hav-
ing had the most time to evolve to take full advantage of Earth's
varied habitats, including the living environments of their fellow
beings.
By trading genes and acquiring new heritable traits, bacteria ex-
pand their genetic capacities-in minutes, or at most hours. A huge
planetary gene pool gives rise to temporarily classifiable bacterial
"typ.r" or "strains," which radically and quickly change, keeping
up with environmental conditions. Bacteria in the water, soil, and
'Whereas your genes
air are like the cells of a growing global being.
are inside a body with a discrete life span, a bacterium takes and
gives out its body's genes in and from the surroundings. Although,
of course, like all life, bacteria can be killed by starvation, heat, salt,
and desiccation, these microbes do not normally die. As long as the
ambience permits, bacteria grow and divide, free of aging. Unlike
the mammalian body which matures and dies, a bacterial body has
no limits. A disequilibrium structure thrown up by an evolving uni-
verse, it is, in principle, immortal. Sequestering order in a disor-
dering universe, the silent bacterial biosphere preceded all plants,
aninrals, fungi, and even the protoctist progenitors of all these forms
of larger life. Without the bacterial biosphere no other life would
ever have evolved, nor would it live today.
Bacteria are the most tenacious beings known. Some survive ex-
treme environments in the dry Sinai Desert, others in the salts of
the Red Sea. Some inhabit Antarctic rocks; others thrive in the Sibe-
rian tundra. More bacteria inhabit your mouth right now, even if
you've just brushed your teeth, than there are people in New York
City.
Bacterial tenaciry should not be underestimated. This entire
planet is bacterial. Human technologies and philosophies are per-
 98 What ls Life?

mutations of the bacteria. Eating, infecting, and irreversibly merg-

ing with one another, bacteria spun offpowerful new prodigies: the
protoctists, fungi, plants, and animals-all of which keep alive the
metabolism and movement of the bacteria from which they derived.
Scientists were originally surprised when they detected hemo-
globin, the red protein pigment in human blood, in legume roots
of pea, bean, and alfalfa plants. Had vegetables somehow appropri-
ated this red, oxygen-carryng iron molecule from the animals which
feed on them? Possibly. But hemoglobin now has been discovered
in the filamentous, sulfur-oxidizing bacterium Vitreoscilla. More
likely, therefore, hemoglobin evolved in the bacterial ancestors of
both plants and animals. Hemoglobin is chemical evidence of a
"blood tie" to early life-a blood tie that evolved long before blood.
Molecules like green ctrlorophyll and red hemoglobin that evolved
in colorful and wily bacteria suggest the extent to which they are
our kin.

FROM PLENTY TO CRISIS

Threatened by the indifference of the matter from which it evolved,

life was enveloped in a world of dangers. At each point in its evo-
lution, life has raised the stakes of existence. Overcoming itself, ex-
panding its sensitivities and capacities, life has plunged into new
realms and new risks-but also new opportunities.
Consider: life produced startling new forms during the time it
was exclusively bacterial. Only recently has science revealed the daz-
zling events that occurred early on in life's evolution. To be recog-
nized and understood by organic beings in the Cenozoic, these most
ancient wonders of natural history had to await micro-, molecular,
and paleobiology. Subvisible metabolic changes and the power of
our planet to process its soil and atmospheric gases can be inferred
today from an arcane fossil record of rnicrobes. And vestiges of the
past are revealed by teasing through molecules and molecular
processes that characterize the biosphere today. Moreover, devel-
 Masters of the Biosphere 99

opments wrought by the early evolution of life did not stay within
the bounds of cell membranes: they became geological, and ulti-
mately planetological. The Archean bacteria changed Earth forever.
Life's tendency to reproduce to the limit produces shortages and
pollution. Responding to changing environments, bacteria fo-
mented a series of "crises." Each crisis was eventually overcome by
evolving many new metabolic pathways, but these, in turn, led to
new shortages, new polluting substances, new dangers for life on
Earth.

BREAI(FAST FERMENT

The original form or forms of cellular, reproducing life spread rap-

idly across a planet whose matter, at first, was little different from
that within the bodies of the biosphere's first beings. The earliest
bacteria grew by fermentation; they broke down organic com-
pounds, sugars, and similar small organic compounds for their chem-
ical energy and food. The biosphere then required no "primary pro-
ducers."The planet was awash in "food" manufactured and renewed
by preliving processes such as solar radiation, the churning of mat-
ter by heat in the deep Earth, and lightning.
Fermenters excrete acids and alcohols, compounds that contain
less energy than the food ingested.Modern fermenting bacteria, for
example, make lactic acid from milk sugar and metabolize gtape and
grain sugar to make the alcohol in whisky and wine. Although the
early Earth had no fruit or indeed plants of any kind, ancestral fer-
menters Gasted on sugar. Laboratory experiments simulating the en-
ergized atmosphere of early Earth have shown that many energy-
rich organic compounds form spontaneously. These compounds
include simple sugars, like the fermentable glucose and the ribose
of RNA. Early life fed on these primeval sweets, replacing carbon-
hydrogen compounds in the environment with its own composi-
tionally similar, but distinctively autopoietic, bodies. There may
be no such thing as a free lunch, but life did, it seems' enjoy the
 100 What ls Life?

cosmic equivalent of a free breakfast. Sugar provided Grmenting

bacteria both with food, for building carbon compounds of self and
progeny, and with energy, for doing the work of maintenance.
One variery of fermenting bacteria thought similar to very early
life is Thermoplasma. These sulfur- and heat-loving bacteria are
grouped with the archaebacteria because of the character of their
RNA. Like all life, Thermoplasmahas a membrane. But unlike other
bacterial cells, except those of mycoplasmas, it lacks a rigid cell wall
outside the membrane. The absence of an outer cell wall gives Ther-
moplasma an ill-defined, mutable shape. In most bacteria the cell wall
is made of sugar-bound peptides, wholly unlike the phospholipid-
protein material of the flexible membrane. Perhaps wall-less bacte-
ria like Thermoplasma evolved from an ancient bacterium prior to
the evolution of any cell walls. Or perhaps the walls were lost. Cer-
tain infectious bacteria that lack cell walls can be perilous to hu-
mans because they resist penicillin and other antibiotics that work
by inhibiting cell wall growth-which is why, at base, antibiotics
cannot harm our animal cells directly. (Our cells have membranes
but no cell walls.)
Wall-less, fermenting life freeloaded, utilizing available sugars and
other energy-rich compounds from the sweet primeval environ-
ment. None of the earliest bacteria were forced to make their own
food. But at some point, the press of expanding generations meant
that free food was decreasing faster than it was restored. A crisis was
inevitable.

GREEN, RED, AND PURPLE BEINGS

Biologists classify as heterotrophs bacteria which, unable to produce

food, obtain their carbon and energy from the outside. We and nearly
all animals are heterotrophs, too. Lacking plastids or symbiotic al-
gae, our bodies do not photosynthesize. We must find food and en-
ergy in the prefabricated form of other living beings, such as rhe
tissues of plants or those derived from plants (animals and fungi). It
 Masters of the Biosphere 101

was thus only a matter of time before the first wave of bacterial het-
erotrophs confronted a biological crisis: environmental food sup-
plies became limited, unpredictable, scarcer and scarcer. Ferment-
ing life could not count on the dwindling sugars of the planetary
pantry. Faced with starvation, some lucky fermenters somewhere
evolved the capability to make their own food, thereby initiating
the great lineage of green and purple beings.
The most important metabolic innovation in the history of the
planet was the evolution of photosynthesis. By way of photosyn-
thesis, life freed itself from energy scarcity; from then on life was
Iimited primarily by the scarciry of one material building block or
another. Photosynthesis appeared in bacteria. Mining the energy in
sunlight, these first food-makers (probably green-colored sulfide
scavengers, like modern Chlorobium) generated food and usable en-
ergy for the rest of the biosphere. They were the first autotrophs.
For life, the most important form of radiation impinging on
Earth's surface is neither short-wave ultraviolet radiation, which de-
stroys biochemical processes, nor low-energy longer wave infrared
radiation, which humans perceive as heat. Rather, life depends on
the medium-wave radiation of visible light. In photosynthesis the
energy of a photon from the sun's visible light excites an electron
in a molecule of chlorophyll, which then transfers the excess en-
ergy to a molecule of ATP. ATP's great contribution to life is that
it allows an organic being to use energy when it is needed-notjust
when the sun (or ingested food) happens to make it available. ATP
is the first-order way that life banks for the future. But ATP as a
tool for energy storage is itself limited. Longer term and higher vol-
ume storage can be achieved by using the ATP to build sugars from
atmospheric carbon dioxide and some source of hydrogen. Permit-
ting cells to make sweets and genes inside themselves, photosynthesis
thus freed life from its early diet of environmental candy.
Today's green sulfur bacteria, such as Chlorobium uinosum, are pho-
tosynthetic. Their ancestors may have been among the first photo-
synthetic beings. Today confined to the bacterial underworld where
 102 What ls Life?

oxygen will not harm them, early on they could have dominated
the surface. The early atmosphere, lacking in oxygen, would not
have bothered them, and it was an extravagant source of carbon
dioxide. The anaerobic metabolism of the green sulfurs thus sug-
gests an ancient heritage.
'Whereas
early fermenters had to eke out a living on ever-scarcer
organic crumbs, early photosynthesizers could literally make them-
selves appear from air. When hydrogen still eisted on Earth as a
free gas, the photosynthesizers had no problem finding the hydro-
gen component for synthesizing sugars. Out of hydrogen gas drawn
from the atmosphere and carbon dioxide drawn from the atmos-
phere the first photosynthesizers crafted their tiny bodies.
Another ready source of hydrogen was hydrogen sulfide. Repro-
ducing and softening once-sterile land, green sulfur bacteria took
in hydrogen sulfide (H2S) spewed from vents and volcanoes at the
ruptured surface. Their waste product was (and is) elemental sulfur
(S); that is why they are called sulfur bacteria. Unlike algae and plants,
which obtain their hydrogen atoms from water (HrO), the green
sulfur bacteria did not expire oxygen gas. Instead, they deposited
elemental sulfur, and did so with abandon. Hydrogen sulfide, ow-
ing to the planet's tectonic restlessness, remained plentiful even af-
ter free hydrogen had escaped from the atmosphere. The hydrogen
sulfide pathway for photosynthesis thus proved to be a fine straregy
for early life.
Scanning the horizon during the Archean eon would have vis-
ited upon the time-traveling eye a quiet riot of glistening color.
Bright red, green, purple, and orange, the photosynthesizers colo-
nized the surface of new volcanic terrain, encroaching upon lava
flows, hardened pumice, and sparkling black sand. Among the fan-
tastically successful bacterial beings were certain heterotrophs that
evolved swimming as a means to obtain new sources of food. Some
of these were red beings that derived their hue from rhodopsin, a
light-sensitive pigment which, like the green chlorophyll pigment,
captures energy for ATP to store-but from a different portion of
 Masters of the Biosphere 103

the spectrum of light.The rhodopsin in today's varieties of halophilic

(salt-loving) archaebacteria exemplify the fundamentally conserva-
tive chemistry of life, as rhodopsin is found in retinal rods of ma-
rine fish and is used in our own vision, especially when light is dim.
The time-traveling eye, moving forward in time somewhat be-
yond the origins of the green sulfurs and the red halophiles, would
eventually spot a new photosynthetic form: purple sulfur bacteria
(see plate 7). Their brilliant innovation was not so much a differ-
ently pigmented approach to photosynthesis but an abrlity to toler-
ate oxygen. Their tolerance was not, however, complete' Purple sul-
fur bacteria living today can tolerate oxygen only in the temporary
darkness of nightfall or other cover. Yet photosynthetic beings, given
their autopoietic imperative, must dwell in sunlight; none can live
for long in the dark.

OXYGEN EXCITEMENT

The purple sulfur bacteria of the Archean that could tolerate oxy-
gen had an advantage. Oxygen was now beginning to creep into
the environment. A new form of water-using bacteria, the cyano-
bacteria, had put it there.
Sometimes still called plants or "blue-green algae," the cyanos are
neither plants nor algae. Cyanobacteria wreaked havoc with the plan-
etary environment-more so than has any life form before or since.
Life had always existed in a medium rich in hydrogen: HzO, water.
Nonetheless, life's supply of hydrogen to make the organic com-
pounds of its bacterial bodies had heretofore come from sugars such
as glucose (C6Hrz06), or from hydrogen and hydrogen sulfide in
the air. The cyanos evolved when photosynthetic bacteria, em-
ploying a unique green chlorophyll system, mutated from purple
predecessors to get their hydrogen atoms from water. Splitting apart
hydrogen dioxide (water) into its constituent atoms, the blue-green
bacteria assembled hydrogen into themselves.
Far more abundant than stinking hydrogen sulfide, clear water
 104 | wr.,ut ts Life?

'Wherever
abounded. they could access water and sunlight, the blue-
green bacteria grew. Today in light and water, these oxygen-mak-
ing photosynthesizers, still capitalizing on their ancient metabolic
innovation, continue to thrive. More than ten thousand kinds have
been cataloged. They are found virtually everywhere-on damp,
dimly lit walls at the mouths of caves, following the slow leaks of
refrigerators, on boat decks, boulders, cliffs, drain pipes, toilet tanks,
and shower curtains. They exist in the Red Sea, in boiling springs,
nuclear reactor cooling tanks, the Sinai Desert, across the Siberian
tundra, and under the Antarctic ice. Cyanos have been cast by some
scientists as the beings most likely to reproduce were they to be
strewn about the red surface and polar dry ice caps of Mars.
The rampant growth of cyanobacteria on Earth was not a local
phenomenon. \X,'herever blue-green cyanobacteria grerv they in-
corporated the "H" from HzO into their tiny bodies and released
into the air the "O" as 02, ox-ygen gas. Highly destructive to all cells
when it causes miniature biological explosions, oxygen gas was fatal
to most forms of early life. Even today it is toxic in high concentra-
tions. Oxygen combines dangerously with enzynres and other pro-
teins, nucleic acids, vitarnins, and lipids. And oxygen produces "free
radicals"-reactive, short-lived chemicals that interfere with meta-
bolic systems. Nutritionists have implicated free radicals in the hu-
man aging process, recommending anti-oxidants such as vitamin E.
In the Archean eon oxygen reacted, sometimes violently, with
atmospheric gases such as hydrogen, ammonia, methane, and hy-
drogen sulfide. Origins-of-life theorists agree that life on Earth has
little to no chance of re-evolving here because free oxlrgen would
oxidize hydrogen-rich chemicals crucial ro srarting any life. The
outer solar system, however, is a different story. Free oxrygen is not
a major constituent of the atmospheres of planets like Jupiter or
moons like Titan. Indeed, if life were ro evolve again it would be
far more likely to do so in the gaseous hydrogen-carbon environ-
ments of the outer solar system, where free oxlrgen is not around
to interrupt early life's orrygen-intolerant chemical systems.
 Masters of the Biosphere 105

Releasing oxygen, the blue-green bacteria enveloped the world

with oxygen atoms. Oxygen accelerated change as it sped up chem-
ical reactions. Crowding out their purple brethren, blue-greens
swarmed the photic zone, the region illuminated and irradiated by
the sun and extending to no more than rwo hundred meters below
the ocean surface. In warmer months cyanobacteria grew most vig-
orously. Sliming along surfaces, trapping and binding sediment, they
formed reefs along the coast and damp, textile-like masses inland.
Rust, iron oxide, formed as bacterial waste oxygen reacted with
dissolved iron. Since iron is Earth's fifth most abundant element,
iron oxides formed in great profusion and, as solids, settled quietly,
year after year, to the bottoms of lakes and seas, including basins re-
cently hollowed out by meteorites. Cyanobacteria flourished in the
heat and grew more slowly in the cool, day after day, summer after
winter, warm millennium after millennium, producing more or less
oxygen in accordance with cyclical and extended climatic changes.
Environmental oscillations led to an alternation berween oxygen-
rich and oxTgen-poor varieties of iron ore; magnetite is less oxi-
dized and hematite more so. The seasonal and climatic shifts in pop-
ulation growth, metabolic activiry and communiry structure of
oxygen-producing bacteria, coupled with changing environmental
conditions, formed great banded rocks. In North America banded
iron formations in ancient, exposed rocks now stretch from eastern
Ontario to the western edge of Lake Superior. They are the source
of the iron used in Detroit-built cars.
TWo thousand million years ago not only iron but sulfur, uranium,
and manganese oxidized all over the world. Exposed to bacterial
waste gas, they became hematite, pyrite, uraninite, and manganese
dioxide. As atmospheric oxygen increased, banded iron, which har-
bors nonoxrygenated minerals, gave way to "red beds"-rust fbr-
mations that formed all over the world. The rock record of oxidized
minerals in Earth's crust testifies to the addition of oxygen to our
planetary atmosphere in a 4oo-million-year stretch from z,zoo to
r,8oo million years ago. Eventually no more minerals were left that
 106 What ls Life?

had not already reacted with oxygen, so the excess gas with no place
left to go began to accumulate in the air.

QUINTESSENTIAL POLLUTERS, QUINTESSENTIAL RECYCLERS

In what passes for humiliry and respect for the ways of nature, mod-
ern humans worry about our pollution of Earth. Pollution is cer-
tainly distressing. But it is hardly unnatural. The pollution crisis
effected by all-natural, blue-green bacteria was much worse than
any we have seen lately. It destabilized the planetary environment.
It made Earth inflammable, and to this day only the ancient surfeit
of oxygen permits us to strike a match to make fire.
Human industry has increased the concentration of ozone-un-
friendly chlorofluorocarbons in the atmosphere some one hundred
times, up to about a billionth of a percent. This degree of change
cannot even begin to compare with the effect upon the global en-
vironment wrought by the blue-greens. By growing, they increased
atmospheric oxygen concentration from less than one part in r,ooo
million to one part in five (zo percent). And Earth's protective, ul-
traviolet-shielding layer of ozone (O3, a three-orTgen molecule) was
built up largely by "al1-natural" pollution in the first place.
But if pollution is natural, so is recycling. Our fresh air is one-
fifth orrygen. Today the ozone layer protects animals such as our-
selves from ultraviolet skin cancer, cataracts, and compromised im-
mune systems. One of the greatest turnarounds in evolution was
the transformation of a once-fatal form of air pollution-oxygen-
into a coveted resource.
Far from destroying the planet, oxTgen energized it. In far-from-
equilibrium systems, waste products necessarily accumulate. But what
may be garbage to one is dinner or building materials for another.
Bacteria, the greatest metabolic innovators, are not only the great-
est polluters but also the greatest cleaner-uppers. Our own chenri-
cal abiliry to use oxygen for energy derives from bacteria. Natural
pollution recycling by bacteria extends to a host of other substances.
 Masters of the Biosphere 107

Green and purple sulfur bacteria, starting with sulfide, produce sul-
fur globules and sulfate (both are more oxidized forms of sulfur),
which suspend or dissolve in sea water. This sulfur is taken up and
recycled by fermenting or sulfate-reducing beings.
Bacteria, in another one of their global megatricks, take nitro-
gen gas lost to the air and return it to all other living beings, where
it is essential for the construction of proteins. Only a few types of
bacteria own this miniaturized industry, as only a few are capable
of breaking the strong triple bonds of molecular nitrogen and then
sequestering the nitrogen atoms into organic molecules without oxy-
gen sneaking in somewhere along the way. Bacteria thus "fix"
gaseous nitrogen-by far the most abundant gas in the atmosphere-
into organic compounds for all the living beings onEarth. Nitrogen-
fixing structures, called heterocysts (large cells in chains mainly made
of smaller ones) were left z,zoo rnillion years ago in the fossil record.
Cyanobacteria with heterocysts can fix N2 gas and make it available
as food (see plate 8).
The creative recycling metabolism of bacteria, combined with
the imperative of autopoiesis, insures the biospheric flow of nitro-
gen, sulfur, carbon, and other compounds. Once nitrogen, for ex-
ample, is fixed into protein and nucleic acid inside bacterial het-
erocysts, and once these proteins make their way through the food
chain (degraded to amino acids and variously rebuilt along the way,
with some leakage into the atmosphere as waste), bacteria are sum-
moned once again to do what only they can do: fix nitrogen back
into organic molecules. The organically bound nitrogen in proteins
and amino acids takes many routes. Some is degraded to ammonia
(NH, by a diversiry of bacteria. Ammonia is oxidized to nitrite
(NO, or nitrate (NO, by still other bacterial specialists. Nitrite
and nitrate, in turn, fertilize the water, letting cyanobacteria and
others grow. Nitrite and nitrate may be "breathed" by some bac-
teria which vent nitrous oxide ("laughing gas") and nitrogen (Nz)
into the air. Nitrogen gas in the atmosphere must then be fixed
again. The complex cycle never ceases. Although no bacteria yet
 108 What ls Life?

degrade the refractory carbon-hydrogen compounds of most plas-

tics, eventually some will evolve and, not limited by food supply,
they will spread like wildfire from landfill to landfill through the
biosphere.

LIVING CARPETS AND CROWING STONES

Like magic carpets in certain remote corners of Earth, "rnicrobial

p2;5"-lgge numbers of interliving bacteria-have the power to
take scientists back in time. Slick and slimy, often stinking of sulfur,
microbial mats preserve the primeval scene of the early Earth be-
fore oxygen surfeited the air. The moist, multicolored mats found
just inland from the sea feel cool beneath bare feet. Save for a sheet
of algae, sand fly egg, occasional gull overhead, or paleobiologist
footprint, nonbacterial traces are rare in microbial mats.
Modern mrcrobial mats and scums are found ever).where in the
world, but only in a few locations are they not obscured by larger
forms of life. At Laguna Figueroa and Guerrero Negro in Mexico's
Baja California, fronting the coastal town of Beaufort in the state
of North Carolina, at Plum Island and Sippewissett in Massachu-
setts, along the edge of the Great Salt Lake in Utah, and at the delta
of the sprawling Ebro River in Spain, mats are conspicuous. Visible
in places too hot, cold, windy, or salry to support larger life, mi-
crobial mats represent what life on a bacterial planet might have
looked like several thousand million years ago.
Merging metabolic talents, a variery of bacterial forms organized
into layers thrive in the mats that they themselves produce. The sun-
loving cyanobacteria dwell in the upper layers, subtly and continu-
ously transforming carbon, nitrogen, sulfur, and phosphorus-
supplying these to their dependents below. Most are multicellular
gliding filaments. Some are single-celled. Together, shaped as threads,
spheres, or branches, the cyanocolonies may form green jelly balls
or scums in the water. Among the photosynthesizers, the most light-
tolerant, desiccation-resistant forms inhabit the upper sheets of mi-
 Masters of the Biosphere 109

crobial mats; the sulfide-using and dim light-seeking members co-

habit the lower layers. The purple sulfur bacteria navigate the mid-
dle realm, balancing a need for sulfide from the nether regions with
a requirement for sunlight from the higher ones.
Lung-like, not only do the gases in the bacterial community move
up and down daily, but so do the constituent guilds. Yehuda Cohen
and his colleagues at the marine station at the Gulf of Eilat in Israel
have determined that when the sun sets the purple layer of sulfide
bacteria moves up a fraction of a centimeter. Deprived of sunlight,
'When
the blue-greens abruptly stop metabolizing. instantly at the
dimmest daylight the blue-greens resume photosynthesis, they be-
gin again to bathe the purple and green bacteria below with their
oxygen waste. The purple layer retreats.
Like a troop of sea turtles on the march-their shells jutting out
of the shallow ocean in Shark Bay, Australia-are the domed, lay-
ered rocks called stromatolites (see plate 9). The American geolo-
gist Charles Walcott (r85o-r927) called the fossil remains of very
similar but very ancient rocks near Albany, NewYork, "cryptozoans"
(from Greek for "hidden animals"). The locals in Saratoga Springs
and around Skidmore College today call these formations "cauli-
flower Limestone." Although Walcott had an inkling that the round
Iimestone "heads" were produced by life, only recently has it become
clear that "cryptozoans" are stromatolites made by vast hordes of
bacteria. They are, in essence, fossilized microbial mats that took the
form of domes rather than the columns, reefs, and pancakes rypical
of the more spectacular ancient stromatolites (see plates roa and rob).
The microbial communities, led by blue-greens, trapped, pre-
cipitated, and bound up calcium carbonate and grains of volcanic
glass before they died. In Australia, where stromatolites are form-
ing today by the work of live bacterial communities, the trapping
and binding can be studied directly. Stromatolites (which sometimes
make use of environmental silica or even iron rather than only car-
bonate for building material) grow a layer at a time, as photosyn-
thesizing bacteria glide past one another, slipping out of their poly-
 110 What ls Life?

saccharide sheaths, which are carbohydrate encasements sindlar in

chemical composition to mucus. The sheaths are sticky and they bind
sand. The live cyanobacteria, gliding toward the sun, leave behind
their sheaths to be colonized by other shelter-seeking microbes.
Tiapping sediment and precipitating carbonate out of water, some
of these complex mat communities solidifii to make living fortresses
against spume and wave. These fortresses thrive, as many kinds of
photosynthetic bacteria support a plethora of camp followers. Spi-
rilla, spirochetes, coccoids, and spore-formers in lively comrnuni-
ties jostle for space, food, and position.
Some fossil stromatolites (such as those in the rocks of the Pon-
gola Group in Africa, the'Warrawoona Group in the Pilbara region
of 'Western Australia, and the Swaziland in South Africa) harbor rni-
croscopic bacterial imprints. These silica stromatolites, containing
black chert, are noteworthy for the microfossils inside that provide
the best evidence for life in the Archean eon.
Thus, among their other accomplishments, bacteria created hard
structures z,ooo million years before the first animal evolved. Stro-
matolite hillocks would have been a common scene in the late
Archean eon. Like miniature cathedrals, they were an early mani-
festation of life's abiliry to manage its excess. A landscape similar to
that seen at Shark Bay has been in continuous existence somewhere
on Earth since life began.
On a global scale the complex tissues of microbial mats-whether
as living carpet or growing stone-may be as important to bios-
pheric functioning as lung and liver are to us. Bacteria took over
the world and still run it, using their decentralized planetary rne-
tabolism and capaciry for worldwide intraspecies gene transfer.

SO, WHAT lS LIFE? Life is bacterial and those organisms that are
not bacteria have evolved from organisms that were. By the end of
the Archean eon every desert was encrusted with rnicrobial mats
and temporary scums; every hot pool, sulfuroLrs or ammoniacal,
 Masters of the Biosphere 1',t1

boasted hoards of colonists and pushy imrnigrants. Over salt grains

and in rusty pools bacteria fabricated glues and precipitated mag-
netite. Clinging to the cold, barren rocks near the poles and slim-
ing over the volcanic rubble in the tropical shallow seas, greening
the Earth, photosynthesizers exuded their wares to hungry oppor-
tunists. The waste of a fermenter became the food of the acid-lov-
ing swimmer, while the fetid breath of a sulfate reducer provided a
precious raw material to green chlorobia or red chromatia. Every
available piece of real estate on this planet was occupied by en-
lightened producer, busy transformer, or arctic explorer. Naturally
selected offspring survived, but only if lent a plasrnid-borne gene
from a comrnunity member. Gene exchanges were indispensable to
those that would rid themselves of environmental toxins: a protein
to be degraded, a poisonous manganese scum, or a threatening cop-
per sheen to be oxidized or reduced. Replicating gene-carrying plas-
mids owned by the biosphere at large, when borrowed and returned
by bacterial metabolic geniuses, alleviated most local environmen-
tal dangers, provided said plasmids could temporarily be incorpo-
rated into the cells of the threatened bacteria. The tiny bodies of
the planetary patina spread to every reach, all microbes reproduc-
ing too rapidly for all offspring to survive in any finite universe. (Jn-
dercover and unwitnessed, life back then was the prodigious prog-
eny of bacteria. It still is.
 PERMANENT MERGERS

I have also seen a sort of animalcule that had the figure of the
river eels: These were in very great plenty, and so small withal
that I deemed 5OO or 5OO of them laid out end to end would
not reach to the length of the full grown eel such as there are in
vinegar. These had a very nimble motion, and bent their bodies
serpentwise, and shot through the stuff as quick as a pike does
through water.
ANTON VAN LEEUWENHOEK

We cannot fathom the marvelous complexity of an organic

being; but on the hypothesis here advanced this complexity
is much increased. Each living creature must be looked at
as a microcosm-a little universe, formed of a host of self-
propagating organisms, inconceivably minute and as numerous
as the stars in heaven.

CHARLES DARWIN

The greatest division is not even between plants and animals,

but within the once-ignored microorganisms-the prokaryotic
Monera and the eukaryotic Protoctista.
STEPHEN JAY COULD

The appearance of these [protoctisu cells a billion-odd years

ago was the second major event in planetary evolution and led
directly, lineage by lineage, to our own complex selves, brain
and all.
LEWIS THOMAS

1',t3
 114 What ls Life?

THE GREAT CELL DIVIDE

Some z,ooo million years ago, probably at many different sites on

Earth, a new kind of cell evolved from bacterial interactions. The
evolution of these complex new cells from integration of bacterial
symbionts prepared the way for liG in the new, Proterozoic eon.
These new cells were ultimately the result of hunger, crowding, and
thirst among teeming bacteria. These new cells were the first pro-
toctists, and their coming brought the kinds of individuality and cell
organization, the kind of sex, and even the kind of mortaliry (pro-
gramned death of the indivtdual) familiar to us as animals. Bacte-
ria merged. Curbing their viciousness and surrendering indepen-
dence, they explored new ways to persist and reproduce.
Our kind of life, that of the nucleated cell, began long before an-
imals. Amid cell gorgings and aborted invasions, merged beings that
infected one another were reinvigorated by the incorporation of their
permanent "disease." The first new kind of cell-the nucleated cell-
evolved by acquisition, not of inherited characteristics but of inher-
ited bacterial symbionts. These new kinds of cells making up the bod-
ies of unicellular protists and multicellular protoctists would eventually
lead to the final three kingdoms of life yet to evolve on Earth: ani-
mals, fungi, and plants. Our protoctistan ancestors were beings so
exceedingly weird that, if informed in detail of their existence, even
the credulous author of a medieval bestiary might pooh-pooh the
tale as the impossible product of a febrile imagination.
Each and every organic being on Earth is made of one of only
two kinds of cells. Our kind-and that of other animals, fungi,
plants, and protoctists-possesses nuclei. The other kind, the bac-
terial cell, has no nucleus. In 1937 Edouard Chatton, a French ma-
rine biologist, named the latter cell rype "procariotique"; the or-
ganisms that possess this cell rype are prokaryotes (pronounced
"pro-CARRY-oats"). All the rest of us are eukaryotes ("you-
CARRY-oats"), made of nucleated cells (fig. 7). So the presence of
a membrane-bounded nucleus defines a cell as "eukaryotic." All
eukaryotes come from protoctists; bacteria don't. The long DNA
 cell membrane

rotary motor

undulipodium

kinetosome

mitochondria

cell membrane

centriole in aster

Flc U RE 7. lllustrated comparison of a prokaryote (a bacterium) at top and a eukaryote

(a nucleated cell) at bottom. All living cells on Earth are either prokaryotes or eukary-
otes. The nonbacterial kingdoms-Protoctista, Fungi, Plantae, and Animalia-all con-
of organisms whose cells are eukaryotic. Eukaryotes evolved symbiotically from
sist
metabolizing, invading, infecting, and cohabiting bacteria.
 116 What ls Life?

molecules, the genes of eukaryotes, are organized within the nu-

cleus into at least tr,vo and as many as several thousand chromosomes
(humans have forty-six). As we shall see, this sequestering of pre-
cious genetic material inside a special membrane and the firm bind-
ing of DNA into a particular sequence in a particular chrornosome
limited the genetic promiscuiry that was and still is accepted prac-
tice in the bacterial realur.
A giraffe is a eukaryotic organism, made of eukaryotic cells. So
is a daisy. And an ameba. The differences in behavior, genetics, or-
ganization, metabolism, and especially structure between prokary-
otes and eukaryotes are far more dramatic than any between plants
and animals. Those differences mark the great cell divide. Prokary-
otes and eukaryotes thus form the rwo "supergroups" of life on
Earth.
All of one supergroup and a good portion of the other inhabit
the microbial realm. Bacteria, the smaller protoctists, and yeasts and
other small fungi are microbes. The eukaryotic cells of protoctists
and fungi are bigger than the prokaryotic cells of bacteria; but, like
any cell, they must be viewed with a rnicroscope. The path of tran-
sition between the two supergroups is obscure. The evolution from
prokaryotes to eukaryotes, from bacteria to protoctists, was a "ry*-
metry break" that catapulted life to a greater level of complexiry
and gave it different potentials and risks. Not just by gradual muta-
tion but suddenly through symbiotic alliance did the first eukary-
otes form.

FIVE KINDS OF BEINGS

The earliest eukaryotic cells, living on their own, were protoctists that
evolved by permanent bacterial merging. Floating or free-swimming,
some went on to become animals, fungi, and plants.
The protoctists are wide-ranging group of obscure beings. To-
a
day an estimated 2Jo,ooo species include tiny amebas and diatoms,
and giant kelps and red seaweeds. lJltimately, this group gave rise
 Permanent Mergers 117

to familiar plants and animals such as palm trees and clams. But even
as recently as a thousand million years ago not a single animal, plant,
or even fungus dwelled on Earth. Biospheric functions were han-
dled entirely by bacteria and protoctists.
The ungainly name "protoctist" was introduced by anEnglish nat-
uralist with an equally unenviable name: Hogg. John Hogg (I8oo-
r86r) set forth his views in an article published in r86r, just before
he died: "On the Distinctions of a Plant and an Animal, and on a
Fourth Kingdom of Nature."1 (His third was the "mineral king-
dom.") Neither Hogg nor anyone else at that time was aware of
prokaryotic and eukaryotic cells. But Hogg saw that many organ-
isms were neither plant nor animal.
Unlike the term protozoa ("first animals"), with its unfortunate
connotation that organisms ranging from foraminifera to slime nets
were somehow animals, protoctist simply means "first beings." Pro-
toctists are neither animals nor necessarily single-celled. But when
they are single-celled-or otherwise tiny-they are called protists.
Because all animals grow from multicelled embryos, there are, by
definition, no single-celled animals. So-called single-celled animals
are really the protists, the smaller protoctists. Hogg suggested "Reg-
num Primogenium" as the name for this primordial kingdom. Its
founding members are now known to have originated prior to plants
and animals, and yet protoctists continue to thrive on Earth today
(see plate r r).
In GermanyErnst Haeckel also argued for a new kingdom. "These
interesting and important organic beings are the primary creatures or
Protista."2 The Monera-bacteria-were part of Haeckel's proposed
Protista. Haeckel, recall, was not persuade d by Lazzaro Spallanzani's
boiling of mutton broth to kill microbes. It seemed clear to him
that primordial beings simpler than anything yet discovered must
exist. An ardent believer both in evolution and in the spontaneous
generation of matter, Haeckel sought "an entirely homogeneous and
structureless substance, a living particle of albumin, capable of nour-
ishment and reproduction."3
 118 What ls Life?

English biologistThomas Henry Huxley (r825-r895) was taken

with Haeckel's notion of a primordial protein globule. Examining
ten-year-old mud samples dredged up from the seabed ofrthe north-
west coast of lreland, Huxley discovered a mysterious white ooze.
'Were these Haeckel's postulated earliest Protista? LJpon examina-

tion, the granular ooze was seen to consist of tiny calcareous plates.
Excited, Huxley wrote to Haeckel that he had encountered the an-
cestral life form. Indeed in the flush of his discovery, Huxley hon-
ored his colleague by namrng the "organisms" after Haeckel. Both
nren delivered the exciting news that Bathybius haeckelii, the great
IJrschleim (primordial goop), had finally been found.
Only later was it realizedthat Bathybius haeckelii was just marine
sediment. The white slime that appeared whenever Huxley doused
the ooze to preserve it was an alcohol precipitate of organic debris
that included jellyfish stingers. Far fiom being our primordial par-
ent, the lJrschleim was not even alive. Nevertheless, Haeckel's con-
cept focused scientific attention on beings that escaped the di-
cho tom ous plant / antnal classifi cati on scheme.
Today the tendency to divide life into animal versus plant remains.
Fungi, if they exist at all in the popular imagination, are a kind of
gray plant. Smaller protists and bacteria-nor quite life in the pop-
ular mind-are ignored or lumped together as "germs." Academia
still departmentalizes liG into botany, the study of plants, and zool-
ogy, the study of animals. Fungi, bacteria, and certain protoctists are
often forced in this scheme to be plants under the jurisdiction of
botanists. This quaint plant-animal split does not reflect evolution.
The ancestors to plants and animals were neither; rather they were
comrnunities-bacteria that merged to form a new kind of cell.
The first essentially modern classification was invented by Her-
bert F. Copeland (r9oz-i968), a biology teacher at Sacramento City
College in California. Copeland argued for four kingdoms: Mo-
nera (bacteria), plants, animals, and protoctists. He placed all fungi
(molds, mushrooms, puffballs, etc., which he called "lnophyta") into
a subdivision of Hogg's Protoctista. His book, The Classifcation of
the Lower Organisms, published at Copeland's own expense by a so-
 Permanent Mergers 119

called vaniry press, was read by almost no one except Cornell Uni-
'Whittaker
versiry ecologist Robert H. (r924-t98o). Whittaker de-
vised the most useful groupings of all when he removed fungi from
his Protista and recognized them as a distinct "fifth kingdom."
From today's vantage, Whittaker's five-kingdom classification
scheme best reflects evolutionary relationships. One of us (Lynn
Marguli$ has collaborated with zoologist Karlene Schwartz of the
Universiry of Massachusetts at Boston to sharpen the blurred
boundaries of Whittaker's protists. The Kingdom Protoctista, which
Whittaker limited to unicellular and the smallest multicellular be-
ings, now includes larger organisms that are not plant, animal, fungi,
or bacteria, such as seaweeds.

TWISTS IN THE TREE OF LIFE

The story of how a human-a being made of nucleated cells-

evolves from an ameboid being-a nucleated cell-is bizarre. But
even this story has a preamble: the evolution of a cell with a nu-
cleus. How did such a cell evolve?
The quick answer is by the merging of different kinds of bacte-
ria. Protoctists evolved through symbiosis; twigs and limbs on the
tree of life not only branched out but grew together and fused. Sym-
biosis refers to an ecological and physical relationship between two
kinds of organisms that is far more intimate than most associations.
In Africa, for example, plovers pluck and eat leeches from the open
mouths of crocodiles without Gar. Bird and beast in this instance
are behavioral symbionts; crocodiles enjoy clean teeth in the com-
pany of well-fed plovers. Bacteria live in the spaces between our
teeth and in our intestines, mites inhabit our eyelashes; all these tiny
beings draw nutriment from our cells or our uneaten food, as cells
are shed or as they excrete organic excess. Symbiosis, like marriage,
means living together for better or worse; but whereas marriage is
berween two different people, symbiosis is benveen two or more
different rypes of live beings.
Organisms form many kinds of symbioses, but the most awe-in-
 120 What ls Life?

spiring is the exceedingly close association known as endosymbio-

sis. This is a relationship in which one being-microbe or larger-
lives not just near (nor even permanendy on) another, but inside it.
In endosymbiosis, organic beings merge. Endosymbiosis is like a
long-lasting sexual encounter except that the participanrs are mem-
bers of different species. Indeed, some endosymbiotic linkages have
become permanent.
Bacteria, masters of symbiosis in general, are also the best en-
dosymbionts for at least four reasons. First, because they have been
entering into stable relationships with one another for over 3,ooo
million years, they are good at forming permanenr relationships. Sec-
ond, their tiny bodies fluidly lose and acquire genes, making them
amenable to rapid genetic change. Third, bacteria have only alim-
ited expression of individualiry; no circulating antibodies guard
them-an "infection," far from being rejected as it might be in an
animal with an immune system, can thus become the basis for life-
long association, a mutual evolution. Fourth, bacteria's vast chem-
ical repertoire leads to a tendency for metabolic complementariry
less often seen in associations between akeady highly individualized
members of plant and animal species. Of course, given time, some
plants and animals may come together as closely as some bacteria
have.
Symbiosis produces new individuals. "'W'e" could not synthesize
B or K vitamins without bacteria in our gut. Cows and termites are
not themselves without the swimming fermenters in their digestive
systems-protists and bacteria that break down grass and wood.
Some algae living inside translucent flatworms are such good
providers that the worms have atrophied mouths; the close-mouthed
green worms "sunbathe" rather than seek food, and the endosym-
biotic algae even recycle the worm's uric acid waste into food.
Thousands of other strange partnerships exist. All of the estimated
20,ooo lichens, for example, began as symbiotic associations of al-
gae with fungi or of cyanobacteria with fungi. But the most im-
portant symbioses were those that led to the eukaryotic ce1l.
 Permanent Mergers I nt

Today most protoctist cells and all plant, animal, and fungal cells
contain mitochondria. The oxygen respiration that keeps members
of the youngest four kingdoms of life alive takes place inside these
particular organelles. (Like organs within bodies, organelles are
functioning structures within eukaryotic cells.) Mitochondrial or-
ganelles look like bacteria. They even grow and divide in rwo at
their own pace within the larger cell. They are thought to corne
from bacteria-but after more than a thousand million years of as-
sociation they cannot survive outside the confines of the cell.
The cells of plants and some protoctists, all algae, also possess col-
orful bodies called plastids. All the photosynthesis undergone by al-
gae and plants happens inside the DNA-containing plastid organelles.
Plastids contain the same pigments and other biochemicals found
in the spherical, oxygen-producing blue-green bacteria that thrive
in the ocean. Coincidence? W'e don't think so. Indeed, DNA in the
plastids of the cells of the red seaweed Porphyridium is closer in its
nucleotide sequence to that of certain cyanobacteria than it is to
DNA in the nucleus of the red seaweed itself.
Such genetic evidence links the cell organelles to their origin from
free-living bacteria in a definitive (and now virtually undisputed)
way. Genetic similarities that cross kingdoms are the biological
equivalent of ancient "fingerprints," proving that photosynthetic or-
ganelles did not evolve gradually by a buildup of mutations in the
DNA of plant and algal progenitors, but suddenly, when digestion-
resistant bacteria took up residence in larger cells. In a moment we
will return to the question of how the bacteria that became mito-
chondria and plastids found their way to their current, cozy loca-
tion inside the cell. But, to be chronologically correct, we must first
explore what may be a still older, and deeper, symbiosis.

SQUIRMERS

Nearly all biologists now accept that certain bacteria, after a period
of chemical negotiation and gene transfer, began as symbionts and
 122 What ls Life?

k*
!s
Ir
t
J

.!1sd
FICURE 8. Trichonympha, a chimerical protist. Phylum: Archaeprotista. Kingdom:
Protoctista. This being, as peculiar structurally as any to be found in a medieval bes-
tiary, is composed of a large protoctist host and a swarm of both undulipodia (its or-
ganelles in the front) and symbiotically attached spirochete bacteria in the rear. Iri
chonympha is itself symbiotic in the termites' hindgut, a microscopic zoo containing
many different sorts of protists and bacteria that together aid in the digestion of wood.

became the mitochondria and plastids of larger cells. Most biolo-

gists, however, reject or are ignorant of another idea. Nonetheless,
circumstantial evidence suggests that a still older bacterial symbio-
sis preceded the acquisition of these organelles.
Before oxygen users infected anaerobic, swimrning protists to
form alliances, and before blue-green bacteria were engulfed by these
alliances, faster bacteria seem to have conjoined. Tiansforming from
free-living bacteria to become parts of cells, wriggling spirochetes
may have conferred their considerable powers of movement upon
the outside, and then the inside, of victims that became ancestral
cells. Today's spirochetes are proton-powered bacteria that ferment
carbohydrates and whip about like possessed corkscrews. The most
rapid swimmers of the entire bacterial kingdom, they literally screw
their way through mud, tissue, and slime.
Thriving in saliva, the crystalline sryles-digestive tissues-of oys-
 Permanent Mergers | 'tZZ

ters, the hindguts of termites, and a thousand other equally inge-

nious niches, invasive spirochetes are one of the most successful life
forms on Earth. And they form alliances-often reversibly attach-
ing to larger organisms and propelling them along. Some protist
cells-such as Mixotricha paradoxa and Tiichonympha-htve gone so
far to evolve holdfast structures where free-living spirochetes are
as
encouraged to reversibly "dock"-with their engines running (fig.
8). The spirochetes actively feed on the metabolic leftovers of the
cells to which they attach. The symbiotic advantage is obvious: the
squirming spirochetes move the cells that feed them. A Mixotricha
or Tiichonympha celfwithout spirochetes is like a boat lacking a mo-
tor or like a teenager without a car. Consortia able to swim quickiy
have more opportunities than their sluggish predecessors to find
food, escape predators, and meet mates. But external spirochetal at-
tachments are not the whole story (fig. 9).

Protoctist cells, huge compared to bacteria, display incessant inter-

nal movement. Bacterial cells, lacking any internal movement and
real chromosomes, do not divide mitotically; they do not perform
"the dance of the chromosomes." Mitosis, the chromosome style
of cell reproduction, is widespread among protoctists and univer-
sal in their animal, plant, and fungal descendants. Matching chro-
mosomes line up and move to opposite poles in a kind of mi-
croballet. At the mitotic poles in animal and many protist cells are
centrioles, structures resembling rotary telephone dials that may be
the remnants of spirochetes that long ago entered larger cells to feed
(fig. ro).
The mitosis by which most eukaryotic cells divide ensures that
chromosomes doubled in the parent are partitioned evenly into fwo
offspring cells. Mitosis seems indispensable as a genetic filing and
distribution system for the huge quantities of DNA that most eu-
karyotic cells contain. In each episode of mitosis a series of tiny pro-
tein tubes, microtubules (collectively called the mitotic spindle), ap-
pears. At the end of the process, when one cell has become Nvo,
 bacteria

\
bacterium

undulipodia

FIcURE 9. Spirochetes attach to other bacteria and eventually

become the undulipodia of larger, now eukaryotic cells.
 early stage
of division

chromosomes
appearing;
nucleolus
disappearing

early stage
of chromosome
separation

reproduced
centrioles

reproduced

FIcURE 10. Differentstagesof mitosis, the usual method of chro-

mosomal separation during cell division, or reproduction, of eu-
karyotic cells. The great internal movement in cells with nuclei,
compared to its absence in bacteria, may result from the 2,0O0-
million-year-old remnants of rapidly squirming spirochetes.
 126 What ls Life?

this mitotic spindle disappears. The chromosomes attached to the

tubules of the spindle line up along the plane of the cell's equaror.
These chromosomes, which doubled earlier, now separate as each
half moves along the spindle to the opposite side of the cell. The
chromosomes now at the poles uncoil as the cell proceeds to divide
into rwo. The enigmatic mitotic spindle then fades back into the in-
visibility from which it emerged (see plate rz).
In many animals the centrioles (the telephone dial-like structures)
move to the edges of the cell, where they become kinetosomes by
growing shafts. In cross section, the shafts show a distincrive "9Q)+2"
pattern: nine sets of rwo tubules arranged near the perimeter of the
circular axis, with one set of two tubules at the center. The kineto-
some and centriole have different names only to distinguish the
shafted &om the shaftless phase of the same organelle. Multiple names
for the same organelle are more accidents of history than nomen-
clatural necessities. Two names were bestowed because the distinct
phases were noticed and named long before any relationship between
them was recognized.
The universaliry of the kinetosome shafts-found in plants, an-
imals, fungi, and protists-is strong evidence for a common origin.
The 9(z)+z symmetry is found, for example, in the cell extensions
of the balance organ of our inner ears and in tails propelling the
swimming protist Euglena. The 9(z)+z arrangement can be seen in
cross section in sperm cells of men. Because of their similariry all
9(z)+z shafts that grow from kinetosomes are best referred to by a
common name.-We call them unduLipodia ("waving feet").
The cell reproduction of mitosis-remarkably sirnilar in some
protoctist and all plant, animal, and fungal cells-musr have evolved
in the oldest of these four kingdoms. The protoctists, from which
emerged plants, animals, and fungi, were the first beings with this
motiiiry apparatus necessary for the reproduction of the new cells
with nuclei. But it seems doubtful that the smaller protoctists in-
vented the undulipodium and internal cell movemenr. Motiliry
rather, we believe was the gift of the oldest and original kingdom
of life.
 Permanent Mergers 127

Tantalizingevidence implicates a bacterial origin of the centriole-

kinetosome organelle. Both DNA and RNA have been reported in
these intracellular structures. DavidLuck andJohn Hall at Rockefeller
Universiry in New York Ciry photographed a peculiar, bacteria-like
DNA in the fwo centriole-kinetosomes of the green alga Chlamy-
domonas. Joel Rosenbaum and his colleagues atYale lJniversiry along
with several other scientists working independently, have not, how-
ever, been able to confirm any centriole-kinetosomal DNA in this
green alga.
Living cells are decorated by undulipodia that go by many differ-
ent names. Undulipodia include all cilia and the "tails" of most
sperm. The single-tailed swimmer in bull semen and the hundred-
tailed sperm released by male fern plants are both examples of 9Q)+z
undulipodia. The immotile cilia remnants in the rod and cone cells
of our retinas, the motile ones of the fallopian tube cells that push
a woman's egg toward the womb, and those rejecting debris in our
windpipes are further examples of undulipodia.
It may be that the spirochetes that symbiotically became un-
dulipodia (involved both as cell tails and chromosome movers) have
become so integrated with their partners that they have dissolved
away to mere traces and genetic shadows of their former selves. Like
an artist whose performance of a difiicult routine seems effortless,
the former spirochete genes may be so deeply implicated in cell func-
tion that today they all but defy detection. Oxford Universiry biol-
ogist David C. Smith likens such symbiotic remains to the smile of
the Cheshire Cat, the fictional feline in Lewis Carroll's Alice in Won-
derland who slowly fades away to become nothing but an enigmatic
grin, floating in midair: "the organism progressively loses pieces of
itself, slowly blending into the general background, its former ex-
istence betrayed by some relic."a
The remaining traces of the bequeathers of motility are fewer
and hazier than those left by the cells which gave the green gift of
photosynthesis and oxygen-bubbling respiration. Motiliry in our
view, was the first endosymbiotic acquisition of the nascent eu-
karyote; losing parts of themselves as they evolved, squirming spiro-
 124 What ls Life?

chetes invaded and animated what were to become nucleated cells.

Today, because of time, the evidence is thin. The photographs have
all but faded and the pages have crumbled. Cell history must be re-
constructed from the faintest clues.
A reason to think that spirochete symbiosis preceded the others
is the recent discovery of many protists that have undulipodia but
lack mitochondria. These air-shunning archaeprotists are poisoned
by oxygen-suggesting they date from a time before ancestral pro-
tists had become symbiotic with the o>'rygen-using bacteria that
evolved into rnitochondria. Mitotic cell division in which chro-
mosomes line up on the spindle is universal in animal, plant, and
fungal cells. Only a few dark-dwelling, oxlzgen-shunning, swimming
"amitochondriates" and their obscure relatives (archaeprotists) show
important variation on the mitosis theme.
The absence of intermediates between bacteria and such seenl-
ingly aberrant protists tells us that evolution from bacteria to nu-
cleated but still anaerobic swimmer probably did not occur by ran-
dom mutation alone. The sudden evolution of cells with nuclei and
9Q)+z swimming organelles is best explained by ancient motiliry
symbiosis. When the close connection of undulipodia and mitotic
apparatus is observed in live anaerobic cells, symbiosis becomes the
most parsimonious of all scientific explanations. lndeed, by com-
parison, mutation explanations for the origin of undulipodia seem
far-fetched.
Consider a very ancient ancestor of one of today's bacterial
denizens of hot springs, Thermoplasma. Imagine that ancestor under
attack by spirochetes. Holding firm, its protective membrane resists
penetration. The spirochetes attach on the outside, establishing as-
sociation, as they feed on Thermoplasma's waste. Eventually some gain
entry and merge with the debilitated Thermoplasma to become its
living oars.
Once inside, the spirochete symbionts extend their motiliry skills
to the internal operations of their would-be victim. A sort of bio-
chemical truce prevails, as both sorts of reproducing partner man-
 Permanent Mergers 129

age to coexist. The nucleus, acting today as a sort of central genetic

government, might have evolved as membrane proliferated to keep
the attacking spirochetes from eating out Thermoplasma's DNA. The
captive spirochetes, still moving, ultimately became movers of chro-
mosomes. Mitosis evolved. Spirochete attachments became centriole-
kinetosomes. Perhaps some of these structures-those that have re-
tained the power to reproduce-still contain DNA.
'Whatever the precise scenario for the acquisition of motiliry and

the sometirnes respiratory and photosynthetic talents of eukaryotes,

symbiosis most assuredly belongs in the narrative. Intimate symbioses
were essential to the evolution of cells.

STRANGE NEW FRUIT

The hypothesis that former spirochetes and thermoplasmas merged

to form swimming protists is under investigation now. These merged
beings nray have been the original members of conGderacies of bac-
teria from which all larger life evolved. But what about other sym-
biotic bacteria? How did they become involved?
Think back to those blue-green, photosynthetic bacteria that pol-
luted Earth with oxygen gas. After reacting to make new minerals
such as sulfate (SO4), magnetite (FezO:), and hematite (Fe:O+) all
over the planet's surface, oxygen waste began to accumulate in the
atmosphere. Newly appeared gaseous oxygen killed off untold
hordes of organic beings. Even today certain kinds of cyanobacte-
ria are sickened by their own oxygen; Phormidium, for example, lives
only in muds near other organisms that can quickly use up the oxy-
gen it produces in would-be fatal concentrations.
Early on, cells evolved a tolerance for oxygen at low concentra-
tions. Many modern prokaryotes still function best at oxygen 1ev-
els of about Io percent-half of the atmosphere's rypical concen-
tration today. Oxygen-tolerant bacteria produce enzymes such as
catalases, peroxidases, and superoxide dismutases that react with the
dangerous gas to produce innocuous organic compounds and wa-
 130 | wr''rut ts Life?

'Without
ter. such chemical buffers the carbon of organic tissue is
scorched, torched, and laid to waste by oxygen.
Nonetheless, the mitochondria of our cells come from bacteria that
neither shunned nor merely tolerated oxygen. The bacteria that
evolved into the matrilineally transmitted mitochondria-only the
ovum bequeaths them to the human embryo-exploited oxygen's
great reactivity. Like nuclear physicists devising a way to power space-
craft by using environmentally hazardous plutonium, the mitochon-
drial ancestors turned an intense danger into a radical opportunity.
In perhaps the greatest example ever of recycling, bacteria em-
ployed reactive oxygen to improve cell processes of energy trans-
formation. Oxidizing the material they produced by trapping the
energy of light, purple photosynthetic bacteria increased their abil-
ity to metabolize ATP, the energy storage compound-the bio-
chemical "coin" used by every cell of every living being. Break-
ing down organic molecules and producing carbon dioxide and
water, bacteria diverted the natural combustion of oxygen to their
own purposes. 'Whereas on average two molecules of ATP are pro-
duced by fermentation of a sugar molecule, with the evolution of
respiration the same sugar molecule was made to yield as many as
thirty-six ATP molecules. The new bacreria-including the an-
cestors of our mitochondria-recouped energy from sugar mole-
cules with over fifteen times the efiiciency of their oxygen-poisoned
predecessors.
That our mitochondria's ancestors were oxygen-respiring purple
bacteria has been shown beyond a doubt by DNA sequencing. Like
a village ransacked by barbarians who ultimately became civilized,
fermenting organisms were attacked by oxygen-using predators that
became mitochondrial laborers.'We suspect the earliest hosts were
Thermoplasma-like archaebacteria (already squirming with spirochete
symbionts), able to withstand heat and acid but not free oxygen.
These consortia evolved into the first protists; their spirochetes had
become undulipodia.The Thermoplasmalineage is implicated in this
major evolutionary event because modern representatives resemble
the nucleocytoplasm portions of eukaryotic cells. Thermoplasma aci-
 PermanentMergers I rfl

near\ universal
dophilum, for example, possesses histone-like proteins
in larger life forms-animals, plants, fungi, and some protoctists-
but lacking in other prokaryotes. The presence of histone proteins
in human chromosomes may be a direct inheritance from the pro-
tists that were inVaded by protomitochondria.
The invaders were probably of the "purple bacterial lineage," as
classified by Carl'Woese. These protomitochondria may have been
similar to modern oxygen-respiring, rod-shaped bacteria such as
Paracoccus denitrifcans. This bacterium contains more than forry en-
zymes in common with human mitochondria. More probably, they
were similar to the respiring Bdellouibrio or Daptobacter-modern
predatory prokaryotes in the habit of attacking and multiplying in-
side larger bacteria. Eventually, the victims explode and.a battalion
of intruders merrily swim out. Daptobacter, Bdellouibrio, and similar
unnamed bacteria are necrobes-beings that live off the death of
others. But even if they began as a parasitic infection, the ancestors
to the mitochondria did not stay that way. Fed and protected in a
living environment, the protomitochondria were better offnot de-
stroying their oxygen-intolerant hosts.
Today, although mitochondria still possess their own DNA and
still reproduce like bacteria, they cannot live on their own. The par-
asitism has become permanent: neither partner can escape, neither
can survive separation. The first protists were thus odd couples, the
results of fusion of rwo, or (in the case of plants), at least three once-
independent beings. But unlike the fire-breathing being of Greek
mythology who has the head of a lioness, the midsection of a goat,
and a dragon's tail, these chimeras were real.
How do predators become symbionts? How does a deadly infec-
tion become a bodily part?
The Korean-American biologist KwangJeon at the LJniversity of
Tennessee has already witnessed such a transformation in the labo-
ratory. The answer is thus less a mystery than before. Jeon's exper-
iments dramatically show how bacteria can change from virulent
pathogens to needed organelles.
Like many of science's most amazing discoveries, Jeon's came
 132 What ls Life?

about accidentally to the prepared mind. To his initial dismay he

found one day that his amebas, which he grew in laboratory dishes,
were sick and dying. Microscopic investigation revealed that each
Amoeba proteus was infected with some r 5o,ooo strange bacteria. All
but a few amebas died. Curious about the moribund survivors, Jeon
injected new, healthy amebas with infectious bacteria taken from
the moribund. Most newly injected amebas died within a few days
although, again, some managed to survive. Those that did repro-
duced more slowly. After some months all the survivors were in-
fected. But these survivors had fewer bacteria inside them than those
which had died.
After growing generation upon generation of infected amebas,
Jeon extracted the nuclei from several. He transplanted these nuclei
into healthy, bacteria-free amebas, whose own nuclei had been rni-
crosurgically removed. The amebas with the transplanted nuclei died
on the third or fourth day-unlessJeon rescued them with a needle-
ful of bacterial "infection." The disease had thus become rhe cure,
A deadly bacterium had become a vital cell part.
Decades laterJeon's infected amebas are alive and well and living
in Knoxville, Tennessee. His experiments have been repeated nu-
merous times, and now he observes that the amebas differ in many
features from their never-infected ancestors. Pathogens have become
symbionts on at least four occasions. Symbionts have become or-
ganelles each time. Invader and invaded merge, evolve into new life
forms. Branches on the tree of life do not always diverge but some-
times come together to produce strange new fruit.

WALLIN'S SYMBIONTS

In tgzT the American biologist Ivan Wallin (r 883-r969) wrote, "It

is a rather startling proposal that bacteria, the organisms which are
popular\ associated with disease, may represent the fundamental
causative factor in the origin of species."s He claimed to have grown
mitochondria outside their animal "host cells." Publicly shouted
 Permanent Mergers 133

down by colleagues, Wallin gave up defense of the bacterial origin

of mitochondria while still in his forties.
'W'allin
was almost certainly mistaken, as no one has ever been able
to grow mitochondria by themselves. Nonetheless, Wallin's theo-
retical assertions were prescient. Plant and animal life, he asserted,
had appeared through what he called "symbionticism" or "the for-
mation of rnicrosymbiotic complexes." He meant new species form
by the permanent acquisition of symbiotic bacteria.
'Wallin
Today has been vindicated. His t9z7 classic book, Sym-
bionticism and the Origin of the Species, was the first systematic de-
scription in English of the importance of symbiosis in cell evolu-
tion. Although heresy only decades ago, contemporary biologists
agree that animals, fungi, and plants evolved from protoctist ances-
tors themselves originating from symbiotic bacterial associations.
The crucial piece of evidence unavailable to Wallin until just be-
fore he died was the discovery that mitochondria and plastids pos-
sess their own DNA. Wallin knew, though, that mitochondrta and
plastids tend to reproduce at different times than do the cells in which
they reside-as if demonstrating a residual impulse of their earlier,
wilder days. Bacterial respirers, like those infecting Jeon's amebas,
allied with nucleated swimmers to form the ameba-like ancestors
of larger life forms: aerobic protists. Combining metabolism and
genes, different lineages of aerobic protists went on to evolve into
animals and fungi.
Algae and plants are a further chapter in the same story. In sub-
sequent symbiotic events, swimming protists that had already fully
integrated with purple bacteria (now mitochondria) came to pos-
sess plastids. How? By indigestion. The resistant green bacteria-
the food-remained alive inside transparent, vegetarian protists. A
continuous supply of photosynthate-food made by the trapped
photosynthetic bacteria-rewarded the protist, which quickly de-
veloped a penchant for sunlit waters. Like small farmers who har-
vest their own gardens rather than shop at a grocery, protists which
incorporated their captives became increasingly self-sufficient. In re-
 't34 What ls Life?

turn for the favor of food, the engulGd photosynthetic bacteria re-
ceived a place to live and rapid free transporration inro the sunlight.
These swimmingprotists, which later evolved into algae, were liv-
ing greenhouses. Would-be food, really endosymbiotic bacteria,
photosynthesized inside the luxury prison of live cells. The origi-
nal undigested food was probably similar to Prochloron This grass-
green bacterium grows in the rear chamber-the cloaca-of cer-
tain kinds of marine creatures known as didemnids or "sea lemons."
Prochloron-|1ke bacteria are a good scientific choice for the plastids
of algal and plant cells. Spherical prochlorons and a rod-shaped (but
similarly grass-green) bacterium called Prochlorothrix make precisely
the same pigments-ctrlorophylls a and b-made by green algae and
plants.
Multitentacled hydras, relatives of jellyfish and coral, are white
but tint green when they possess symbiotic green photosynthetic
microbes. Th e snatl, Plachobranchus has gardenlike rows of green plas-
tids under its parapodial folds, part of the digestive tract. The gi-
ant clam, Tiidacna, hosts green dinomastigote algae. Many organ-
isms have allied with photosynthetic bacteria or algae. History
repeats itself.
Grass-green and blue-green bacteria are independent versions of
the plastids of algal and plant cells. Algal plastids need not be green.
Plastids of the alga responsible for the red tint of alpine snow patches
in the late spring and summer ("watermelon snow") are red (see
plates r3a, r3b, and r3c). And in tnzania's Lake Natrum swoop
great flocks of pink flamingos. Red photosynthetic bacreria and al-
gae with red plastids, pigmented with the same carotenoids that color
carrots, grow in the lake. Flamingos look pink because the pigments
at the microbial base of the food chain wind up coloring the bod-
ies of these intriguing birds.
Genetic evidence, DNA, RNA, and protein sequence informa-
tion link red algal plastids to certain cyanobacteria with the same
forensic accuracy admissible in court to convict a rapist whose DNA
matches that of a sperm sample. The multicolored bacteria of the
 Permanent Mergers 135

Archean eon have not gone away. They have joined with other cells
to become the sea-green chloroplasts of garden cucumbers. Oth-
ers have become the brown phaeoplasts of kelp in coastal waters.
Still others lurk today as the red rhodoplasts of dulse, a form of sea
lettuce. If food crops are grown in orbit, on Mars, or on other plan-
ets greened with life, it will be a transhuman phenomenon, part of
the same bacterial expansion that began more than 3,ooo million
years ago on the Archean shores.

MULTICELLULARITY AND PROGRAMMED DEATH

Plants and animals are so complex that it is easy to forget their

original status as colonies of hybrids. Occasionally, however, we
are reminded of our multicellularity. "Hel.a" cells-from the cer-
vix of Henrietta Lane, a woman who lived in Washington, D.C.-
continue to be grown in laboratories around the world, despite
Lane's death from cancer of that same cervix in the r95os. This
morbid medical fact demonstrates our colonial nature as huge col-
lections of nucleated cells organized into tissues.
By symbiosis different varieties of bacteria came together and
made cells with nuclei. These cells with nuclei often cloned them-
selves into multiple copies that stayed in physical contact after re-
production. A Paramecium or Euglena is an "individual" nucleated
cell, already fascinating in its mixture of living beings. But plant,
animal, and fungal life greatly expanded the complexity of the free-
living protist cell by repeating it to make multicellular copies that
ultimately evolved into separate tissues, such as reproductive and
nerve tissue, with distinct functions.
The olfspring of some of these protists, starting in earnest per-
haps a thousand million years ago, failed to separate after they re-
produced by cell division. They began permuting themselves into
colonies, some of whose members died each generation. Thus cer-
tain colonial protists became physically large members of the group,
and the diversiry of protoctists evolved. Looking at modern pro-
 136 What ls Life?

toctists suggests how such colonies could have formed from indi-
vidual cells. Animals, including of course ourselves, are transformed
colonies of protist cells.
Charles Darwin emphasized that evolution occurs as different in-
dividuals pass on their traits by out-reproducing others. But indi-
vidualiry always in flux, form and inreract in a wide
is relative. Cells
array of configurations. Together they form individuals at various
size levels and degrees of interdependence. The alga Chlamydomonas,
with its large green single chloroplast, is a bacterial composite. Voluox,
a spherical confederacy of Chlamydomonas-7lke protist cells, is a green
multicellular descendant of Chlamydomonas, jtst as animals are mul-
ticellular descendants of swimming protists (see plate r4).
The origin of any "individual" large organic being depends on
integrative gene-transferring processes not easily reversed. These
integrative processes first stabilized as the colonial protoctists
evolved from free-living protists. Voluox algae,like other protocrists,
fungi, plants, and animals (but unlike bacteria), do not casually trade
their genes. Larger organisms simply cannot trade genes the way bac-
teria do.
Any single protoctist, plant, fungus, or animal is a member of a
species. Most likely, protoctists were the first organic beings to
form
species and the first whose species went extinct. The origin of in-
dividuals who all belong to the same species is identical to the ori-
gin of the first protoctists. Canadian microbiologist Sorin Sonea
makes a good point when he claims that bacteria, because on a plan-
etary scale they reversibly trade genes, do not have true species.
Species are groups whose members interbreed. A11 bacteria on the
planet can, in principle, interbreed. If anything, they might be said
to form a single, global species.
Species demarcation is thus much more applicable to the pro-
toctists, in which, indeed, it first appeared; so did sexualiry-of the
"meiotic" kind. Fatefully for the future history of life forms such
as ourselves, in protoctists sexualiry became inextricably linked to
death. Bacteria can be killed but they do not naturally die. Certain
 Permanent Mergers 137

protoctists, notably ciliates and slime molds, unlike bacteria, will age
even if external conditions are suitable for health. Aging and death,
in which living cells disintegrate with predictable timing, first
evolved in sexual protoctists. "Programrned" death as the final stop
of a lifelong metabolism was absent at the origin of life-and for a
very long time afterward.
Unlike us, bacteria are imrnortal; they will live until external con-
ditions prevent autopoiesis. By contrast, like us, many protoctists age
and die at the end of a regular interval. Aging and dying is an in-
ternal process called apoptosis or thanosis in technicaljargon. Apop-
tosis arose in our nicrobial ancestors at some time during the evo-
lution of sexual individuals. Strange to say, death itself evolved.
Indeed, it was the fi156-2nd is still the most serious-sexually trans-
nritted "disease."

SEXUAL GENESIS IN THE MICROWORLD,

OR WHEN EATING WAS SEX

Animal sex always involves meiosis. In meiosis, as in mitosis, chro-

mosomes attach to spindle microtubules and are partitioned out to
offspring cells. However, meiosis omits a crucial step: the doubling
of the chromosomes.
Meiotic cell division thus produces rwo offspring cells each with
only half the number of chromosomes that were present in the orig-
inal parent cell. For example, after meiosis a human cell with the
standard allotment of forry-six chromosomes becomes an ovum or
spernl cell with only rwenry-three, ready to find its "other half."
Meiosis, which halves the number of chromosomes per cell, and
fertilization (after sexual intercourse, flower pollination, protoctist
or fungal conjugation), which doubles the number, must go hand
in hand.
Meiotic sex evolved in mitotic protoctists long before any animal
appeared in the record of life. Nevertheless, some modern protoc-
tists show neither mitosis nor meiosis. The giant freshwater ameba,
 138 What ls Life?

.l
,: 1

FIGURE 11. Naegleria, a protist. Phylum: Zoomastigota. Kingdom: Pro-

toctista. The Naegleria amebas caught in the attempt to eat their con-
specific neighbors are shown in this drawing. ln evolution, ingestion fol-
lowed not by digestion but by continued internal existence of the
devoured being was an important means of starting up cellular symbio-
sis. When protists of the same species devoured but did not digest each
other, they sometimes merged nuclei and chromosomes, a commingling
equivalent to the first act of fertilization or mating.

Pelomyxa pdlustris, for example, reproduces by pinching in half its

many nuclei as it pinches in half its body-a single huge cell. Di-
nomastigotes (so-called dinoflagellates) undergo a unique variation
on the theme of mitosis: their DNA is not coated in histone pro-
tein and their chromosomes, unlike those of other microbes, re-
 Permanent Mergers 139

main visible throughout a cell division in which the nuclear mem-

brane remains intact.
Meiosis is a variation on the theme of mitosis. Meiosis likely
evolved in doubled cells that had already divided by mitosis. The
first fertilization event probably satisfied an urge not to merge but
to eat. This could have happened if protist cannibals ate one an-
other. Microscopists sometimes witness microbial wranglings in
which a hungry cell engulfs a neighbor (fig. rr). But the cells do
not always digest what they engulf.
Harvard Universiry biologist Lemuel Roscoe Cleveland (r898-
r97r) saw cannibalized protists live on although half-devoured. The
protists he studied, covered by 9Q)+z undulipodia, are called hyper-
masrigotes. Normal hypermastigotes, which contain only a single set
of chromosomes, live in the swollen hindguts of wood-eating ter-
mites and cockroaches. Cleveland saw the hypermastigotes engulf
one another. He noted that once they did, their membranes merged
to make them into doubled cells. Most of the doubles died. But
Cleveland also saw that some of these doubled beings reproduced.
Although sloppily, a doubled microbial monster would undergo cell
division and give rise to another doubled microbial monster.
Cleveland saw how thwarted cannibalism could have led to the first
setsof doubled chromosomes. Moreover, abnormal cell divisions-
a precursor to the meiosis that takes place in our own cells-could
restore the merged would-be cannibals to their original single set
of chromosomes. Such potential steps in the evolution of sex oc-
curred in a few hours in the protists of Cleveland's laboratory. In
nature, unseen and long ago, they must have occurred spontaneously
many more times. Today when certain hypermastigotes fuse they
form hard-walled resistant structures, cysts, capable of withstand-
ing deprivation. This double form-perhaps originally deriving from
cannibalism-would have protected the protosexual beings engag-
ing in it.
Once upon a time, we think, eating and mating were the same.
Terminal microbial indigestion may seem rather unromantic as the
source of the human sex drive. But Cleveland's picture of hungry,
 14O ] wr,ut rs Life?

serendipitously mating hypermastigotes presents a mix of comedy

and terror appropriate for the origin of sex. In times of scarciry our
unicellular ur-parents would have desperately fed on one another.
Sometimes their membranes would have merged. Doubled, partly
doubled, and huddled in their walled cysts, they waited-abnormal
beings, some now with faulry sets of chromosomes, many of which
would die. Those returning to the original one-set state would be
naturally selected; only they could begin to reproduce normally
again. The doubled and other monsters would tend to die. All the
same, abortive cannibalism and other fusions would often afford pro-
tection against seasonally scanty food and periods of desiccation. [Jn-
able to enter the sturdy quiescence of a sex-induced cyst phase, those
reluctant to eat their kin starved and died of thirst.
The cells in our own animal bodies are in a diploid, or double
chromosome, state except for the protist-like ova and sperm, which
are in a haploid, single-chromosome state. Each animal body is a
sort of diploid husk, morbidly discarded by those haploid sex cells
that manage to produce each generation a fresh new body and thus
continue beyond the death of the "individual." The diploid body
pays the ultimate price-death-for transmission of haploid sex
cells.
Perhaps originally cannibals in distress, chromosomally doubled
protists are our ancestors. Humans and all animals inherited death
from these early eukaryotes. Each generation starts where the last
left offand, depending on who survives, each follows a slightly differ-
ent course. Over time, this leads to new species.
Aggregated into colonies, layers of eukaryotic cells eventually
evolved into tissues. It is an arnazing fact that all zygotes-fertilized
egg cells that survive to become embryos-grow into plants or an-
imals composed notjust of many cells but of several (or many) kinds
of cells that assemble into distinct Although far more im-
tissues.
pressive in animals and plants, protoctists too may be composed of
several sorts of cells showing division of labor. Protoctists recognize
one another; when deprived of water or food, member cells of the
 Permanent Mergers 141

same species can congregate. Thke slime molds: these bacteria-eat-

ing amebas exchange metabolic information, seek out their own
kind.
When food is aplenry the amebas feed alone. But when food is
depleted, each hungry cell secretes a compound that is mutually at-
tractive. Amebas move in the direction of highest concentration.
Together the cells merge to form a moving "shmoo" that grows up-
ward in a slimy mass until its "head" bursts and releases sturdy-walled
cysts, safely transportable by wind and water. A new generation of
amebas will begin if the cysts land in a clement environment.

THE POWER OF SLIME

A focus on our protoctistan origins is humbling. There is no deny-

ing our kinship to this sort of living matter. Humans are integrated
colonies of ameboid beings, just as ameboid beings-protoctists-
are integrated colonies of bacteria. Like it or not, we come from
slime.
An estimated quarter of a million protoctist species dwell in lakes,
rivers, waterfalls, warm springs, damp soils, transient puddles, dew,
and frost, and on the walls of swimming pools and pipes. Plants, an-
imals, and fungi-the visible kingdoms-began as mere fiefcloms
of the Kingdom Protoctista. The initial members of this kingdom
had already evolved a thousand million years before spinning offthe
remaining three kingdoms of life.
Today crystalline beauties such as diatoms, foraminifera, and ra-
diolaria inhabit Earth's oceans. Because protoctists live most abun-
dantly in the tropics, surely far more species are unknown than
known. The best studied include a few notorious killers, such as the
trypanosomes that cause sleeping sickness, Chagas' disease, or leish-
maniasis. In the latter, a disfiguring disease of the tropics, growing,
swimrning protists may devour embryonic mucous membranes ir
utero, caosinghuman babies to be born without mouth or nose. But
together most protoctists cradle the planet in their gentle biochem-
 142 What ls Life?

ical grip. They carpet the ocean with food and supply its oxygen,
churn soil, clear surfaces of bacteria. They cycle sulfur, phospho-
rus, silica, and carbon on a global scale.
Protoctists partake of planetary physiology. Among the most nu-
merous are the coccolithophorids. Though microscopic, these single-
celled algae are one of the few forms of life distinctly visible by satel-
lite. A "bloom" of coccolithophorids may whiten the green warers
oft^the coast of Europe for rwo hundred kilometers.
Not until the ocean water is concentrated by centrifuge in the
laboratory and the sedimented sample magnified some ren rhou-
sand times does the source of the white patch reveal itselfi cocco-
lithophorid scales. Each coccolithophorid is patterned with hundreds
of chalky "buttons." The scales, and the spaces berween them, may
be the natural equivalent of Venetian blinds, serving to provide the
alga's plastids with an optimal quantity of sunlight. The scales re-
leased by dying microbes-rnillions of buttons per milliliter-
change the water to a milky color seen better by satellite than by
ship.
Because salt accumulates in its cell and can destroy it, a cocco-
lithophorid must make complex sulfur compounds that balance in-
ternal ion concentrations. These sulfur compounds are unstable; they
break down to form dimethyl sulfide, a waste gas wafted into the
air. Once released, dimethyl sulfide gas reacts with oxlrgen to pro-
duce tiny aerosol particles of sulfate. These particles seem to be in-
volved in the formation of cloud cover by serving as nuclei for the
condensation of water vapor. Since the cloud cover radiates back to
space and leads to cooler temperatures, a bloom of coccolithophorids
such as Emiliania huxleyi may act as a global air conditioner.
Vast amounts of materials flow through the bodies of cocco-
lithophorids and other photosynthesizing protoctists. Protoctists, not
plants, serve as the base of the entire marine food chain. Floating
protoctists attend to the needs of marine ecosystems far offshore.
Substrate-dependent species support the bulk of communities near
shore.
 Permanent Mergers I raa

FIcURE 12. Stephanodrscus, a diatom. Phylum: Bacillariophyta. Kingdom: Protoc-

tista. Radially symmetrical pillbox diatom. Diatoms, usually tan or brown in color and
prevalent in oceans, deplete water of silica as they make their gorgeous microshells.

Building their skeletons of chalk, glass, organic fiber, and even

exotic salts like strontium or barium sulfate, some protoctists mine
the oceans for trace chemicals. They transform seascapes when, af-
ter building up their bodies' hard parts, they die in droves. Diatoms
deplete silica from the ocean on a global scale to create their ex-
quisite pillbox forms (fig. rz). Radiolaria form opaline shells that
fall like rain through the water colunm, making a film that later hard-
ens into a flintlike sedimentary rock called radiolarite. Foraminifera
make up part of the limestone rock composing the great pyramids
of Egypt.
Protoctists are cosmopolites, infiltrating themselves into the great
 '144 What ls Life?

water and soil mass of Earth. Like the Sphinx, protoctists are re-
combined, fused beings. The first organic beings to form sexually
reproducing species, their cellular quirks are at the core of human
sexuality. The protoctists bequeathed to all subsequent kingdoms of
life the physiological necessiry of death. They, along with bacteria,
are the supreme architects of the living globat environment.

so, WHAT ls LIFE? LiG is the strange new fruit of individuals

evolved by symbiosis. Swimming, conjugating, bargaining, and
dominating, bacteria living in intimate associations during the Pro-
terozoic gave rise to myriad chimeras, mixed beings, of which we
represent a tiny fraction of an expanding progeny. Through corpo-
real mergers disparate beings invented meiotic sex, programmed
death, and complex multicellulariry.
Life is an extension of being into the next generation, the next
species. It is the ingenuiry to make the most of contingency-to
make animals, for example, out of a botched attempt at cannibal-
ism. Life is bigger than the cell, the organism. It includes the bio-
sphere, the surface planetary environment as a whole-&om the for-
mation of marine clouds to the control of ocean chemtstry by
protoctists and their progenitors.
Berween these two realms
lies all of life as we know it.

PLATE 1A. Earth in space.

PLATE 1 B. Mycoplasma. Phylum:Aphragmabac-

teria. Kingdom: Bacteria (Monera). These are some
of the smallest bacteria, with cells less than 0.5 mi-
crometer in diameter.
PLATE 2. Pachnoda, a beetle. Phylum: Mandibulata. Kingdom: Animalia. This
close-up of the intestine of a beetle larva shows a tree-like organ that houses
methanogenic bacteria. Living in the intestine for thousands of generations,
the methanogens have not only found a home, but they have induced devel_
opment of this symbiotic "beetle" organ.
PLATE 3. NASA x-ray photograph of the sun. According to
Russian scientist Vladimir Vernadsky, life on Earth is a mate-
rial system in which stellar energies become transformed into
living ones. Life is not only a global but a planetary-solar
system phenomenon.
PLATE 4. Cylinders of phospholipids produced through a
hydration/dehydration process by David Deamer in his labora-
tory. As seen here through a light microscope at high mag-
nification, they eventually break apart to form liposomes. Such
precellular organic formations were commonplace in Archean
times. The early Earth was like a giant laboratory in which count-
less experiments occurred all the time. ,,Cod," jests organic
chemist Cyril Ponnamperuma, "is an organic chemist.,,
PLATE 5A. Proteusmirabills, abacterium. Phy-
lum: Omnibacteria. Kingdom: Bacteria (Mo-
nera). Nested patterns of concentric terraces
form with the bacteria's repeated cycles of
growth and migration over the petri plate's agar
surface. The living geometry of this photograph
is reminiscent of the nonliving dissipative struc-
tures of plate 5b.

PLATE 58. Autocatalytic chemical reactions of

the same kind shown in plate 5a but using dif-
ferent substances as recursive structures are
thought to give rise to life. This particular
"chemical clock" is a dissipative structure in a
Belousov-Zhabotinsky reaction. The increase in
complexity over time is reminiscent of life. By
reproducing, however, life has increased its
complexity not for several minutes but for sev-
eral thousand million years.
PLATE 5. Myxococcus, multicellular bacterium. phylum: Myxobac-
teria. Kingdom: Bacteria (Monera). These bacteria aggregate into
"trees," up to 1 millimeter tall, when n utrients or water are depleted.
Contrary to common opinion, multicellularity is not a feature
solely of larger organisms but is already an entrenched feature of
prokaryotes.
i* #
j"L E

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*,
ilt'*.
PLATE 7. Chromatium vinosum, purple sulfur bacteria. Phylum: Proteobacteria
(purple bacteria). Kingdom: Bacteria (Monera). These photosynthetic microbeings-
up to 5 micrometers long and from 0.5 to 1 millimeter wide-photosynthesized in
the light long before plants evolved. The purple inclusions are the thylakoid mem-
branes of photosynthetic pigments and enzymes; the spheres are sulfur. These be-
ings, which tolerate oxygen only in the dark, testify to the fact that photosynthe-
sis is an anaerobic process that evolved long before oxygen was present in the air.
PLATE 8. Fischerella, cyanobacterium. Phylum: Cyanobacteria. King-
dom: Bacteria (Monera). An example of bacterial metabolic "supe-
riority," this cyanobacterium fixes atmospheric nitrogen in its hetero-
cysts (each about 1 micrometer wide), making protein. The biochemical
and metabolic repertoire of bacteria makes them crucial to biological
functioning on a global scale.
PLATE 9. Living bacterial "skyscrapers," rocks called stroma-
tolites, in Shark Bay, Australia. Over time some types of mi-
crobial mats are thought to give rise to these strange domes.
The photograph displays live stromatolites, replete with grow-
ing bacteria. Such structures, Senerally much smaller and less
conspicuous, are known from seaside locations around the
world. They occur in both fossil and live form. Huge stroma-
tolites, remnant bacterial landscapes, are common in the rock
record. They represent souvenirs of our bacteria-dominated
planet prior to the evolution of animals, fungi, or plants.

PLATES 1oA AND 1oB. A petrified fossil stromatolite from

Warrawoona, South Africa, is compared with a cut section of
a living microbial mat from Matanzas, Cuba.
P .
LATE 1 'l Mesodinium rubrum (now called Myrionecta rubra) a protist. phy-
,
lum: Ciliophora. Kingdom: Protoctista. This fast-swimming but photosynthetic mi-
croscopic being that dwells in brackish water near the Baltic Sea has relatives jn
Arctrc and Antarctic waters. As neither plant nor animal, it exemplifies organisms
confounding older, two-kingdom classification systems. The reddish hue of the in-
terior derives from diminished symbiotic algae.
 ._*,
Y{:: i
H. ,#!tr

i::r :-::- =::i'

Y:;'- .ia;. .r'' -,1... ,'='li...
' :--+
'j+r<- {=:i'i:::#

PLATE 12. Haemanthussp., African blood lily Phylum: Anthophyta(Angiosper-

mophyta). Kingdom: Plantae. Telophase in the mitotic cells deep inside the
flower. Telophase is a late stage of mltosls-the kind of reproduction of cells by
division that typifres the nucleated cells of all animals, plants, and fungi, and most
protoctists. In mitosis the chromosomes double first and then divide and sepa-
rate into these two masses that become the nuclei of the resulting offspring cells.
ln the mitotic cell division shown here the flower's chromosomes are stained red.
 \;--t\ -1'
, \\-.-\g;,

PLATE '13A. Chlamydomonas nivalis, an alga. phylum: Chlorophyta (green

algae). Kingdom: Protoctista. Snow algae, as seen in this photograph from
Antarctica, have red pigment that masks the green chlorophyll. DNA stud_
ies of the green algae and plants and of marine red algae point to a symbi_
otic origin of the colored cell parts from cyanobacteria. Cyanobacteria, team-
ing up with larger cells, eventually evolved lnto the plastids of all ,,higher,,
(larger) photosynthetic beings, from seaweeds to maple trees.

PLATE 138. Medrum-range photograph of the snow algae Chloromonas

sp. and filamentous fungi. The pigments of these photosynthetic creatures
give the mountainous snow its orange tint.

PLATE 13C. Microscopic view of red snow alga Chtamydomonas nivalis,

so-called watermelon or candy-colored snow. The red carotenoid pigmen_
tation network serves as a photoprotectant from bright sunlight. The cells
are four hundred tjmes smaller than they appear rn this photograph.

r;I: I

E
PLATE 14. Colonies of Volvox. Phylum: Chlorophyta (green algae).
Kingdom: Protoctista. lndividual cells of this green colonial alga resemble
free-living cells of Chlamydomonas. The evolutionary move from unicel-
lular to multicellular " individuality" is a crucial one that has occurred many
times. lt may be happening again as electronically communicating, tech-
nologically interacting human beings form networks required for survival.
 4*-

PLATE 't 5. Lima scabra, a scallop. Phylum: Mollusca. Kingdom: Animalia. A pic-
ture of an adult showing the soft parts of the animal. This bivalve mollusk devel-
ops from a planktic larva (a ciliated trochophore larva) that develops in turn from
a blastula. A blastula type of embryo is a defining characteristic of all animal life.
ln spite of our experience on land, most animal phyla have oceangoing members
such as this one that dwells off the shore of Puerto Rico. The first soft-bodied an-
imals are thought to have evolved in the oceans over six hundred million years ago.
PLATE 15. Drosophilamelanogaster, afruitfly.Phylum: Mandibulata.Kingdom:Ani-
malia. Shown here is an embryo, further developed after the blastula, prepared so that

a red dye tracks the presence of the nervous system in the young animal. Because
fruit flies can be grown (in pint milk bottles in the laboratory) from egg to adult to
egg again in just a few weeks, more is known about the genes and chromosomes, the
growth of the nervous system, the muscles and hormones, the sense organs, the mat-
ing behavior, and all other aspects of the biology of this insect than for any other ani-
mal, including humans. The fruit fly is part of the varied and successful arthropod
group, which includes not only all insects and spiders but also crustaceans such as
crabs and lobsters.
PLATE 17. Undulipodium in cross section. The shaft (axoneme) displays the9(2)+2
arrangement of microtubules. This distinct intracellular organization is found in the
sperm cells of widely diverse beings throughout the natu ral world, f rom men to gin kgo
trees. Electron micrographs of cuts through the shafts of the cilia propelling swimming
paramecia and trichomonads and the cilia that push the egg through a woman's fal-
lopian tube also reveal thtsg(2)+2 pattern. All undulipodia are 0.25 micrometers wide;
lengths vary from less than 1 to over 3,0O0 micrometers (3 millimeters).
PLATE 18A. An angler fish with bioluminescent spots and ribs.
Phylum: Craniata. Kingdom: Animalia. The family (ceratioids) to
which this fish belongs has bioluminescent members that culture
pure strains of glow-in-the-dark marine vibrio bacteria in their
bodies. These deep-sea fish use their bioluminescent symbiotic
organ to lure potential prey, which mistake the protruding ap-
pendage for a small edible fish.

PLATE 188. Photobacterium fischeri, a bacterium. Phylum: Pro-

teobacteria. Kingdom: Bacteria (Monera). This petri plate shows
colonies of bioluminescent bacteria. Many types of fish cultivate
symbiotic bacteria in special organs and put their light to good
use in predator defense, food illumination, or mate signaling.
PLATE 19A. Russula paludosa. Phylum: Basidiomycota. Kingdom:
Fungi. This relatively common forest mushroom is connected to the roots
of nearby trees with which it lives symbiotically.

PLATE 198. Schizophyllum commune. Phylum: Basidiomycota. King-

dom: Fungi. The basidia of this mushroom are borne on the white
double Iines of the gills shown in the photograph.
 PLATE 20. A chloroplast, the intracellular
structure which carries on photosynthesis.
This close-up was taken with an electron
microscope; the organelle is about 1 mi-
crometer long and 0.7 micrometer wide.
A hallmark of plant life, chloroplasts, as has
recently been "proven" by genetic com-
parison, evolved from cyanobacteria that
greened the world long before the origin
of plant life proper. Algae and plants all
seem to have evolved after larger cells
merged with smaller ones. The large cells
fed on but ultimately failed to digest the
once f ree-living cyanobacteria.

PLATE 21. Populus tremuloides, quaking aspen. Phylum: Anthophyta

(Angiospermophyta). Kingdom' Plantae. A stand of quaking aspen in the
San Juan Mountains, Colorado. A similar stand in Utah has been nominated
the largest "organism" on Earth, with each genetically identical tree con-
sidered the stem of a many-treed body. The aspens in the picture, turning
color simultaneously, are not as extensive as the 43-hectare, 6-million-
kilogram stand sharing a single root system in Utah.
PLATE 22. Navicula cuspidata, a diatom alga. Phylum: Bacillario-
phyta. Kingdom: Protoctista. A diatom undergoing meiosis and
gamete formation. Algae, like all protoctists, dwell in water. Plants
eventually escaped water to venture onto land by evolving water-
proofing, such as waxy cuticles, and structural support, such as lignin.
 ::.i

i*$*:

t-: \

\6_.

*F:!.i
. ..r-l'i
r* .,: '''/

PLATE 2 3. Clossopteris scuturn, fossil seed fern. Clossopteris is one of

the extinct groLrps of cycadofilicales whose trees made up the forests over
2)5 ml)tot years ago, before the evo ution of the first dinosaurs. These
anc ent lorests, crushed beneath the surface by {loating tectonic plates,
bccaTrrt' c la .
PLATE 24. Pollen tubes containing sperm nuclei germinate and grow down to mate
with the female egg nucleus deep inside the "embryo sac" of the flower in the dou-
ble mating event that occurs in all angiosperms: one sperm nucleus mates with one
egg, and two other sperm nuclei mate with the egg's "sister nucleus" to form a tissue
that feeds the growing embryo. Here the pollen grains with cytoplasm and nucleus
are stained orange and the pollen tubes are light colored. The pollen grains (male)
and the embryo sac (female) are the gametophyte generations of angiosperms, and
therefore before fertilization all their nuclei have one set of chromosomes. The bio-
logical term for having one set of chromosomes is "haploid." ln the human body
only sperm and egg cells are haploid; plants, however, may produce entire bodies-
whole plants of their life cycle-that are haploid.
PLATE 25. Papaver somniferum, apoppy. Phylum: Angiospermophyta. King-
dom: Plantae. The name of the common poppy means "sleep-bringing," from
the effects of its acrid and narcotic juice. Angiosperm plants, whose evolution
dovetailed with that of mammals, still cast their floral spells on human eaters,
drinkers, and lovers. E. O. Wilson's biophilia theory suggests that we have ge-
netically embedded patterns of emotional response to other life forms. The
color, smells, and tastes emitted by flowering plants captivate with all the power
of their 100-million-year aesthetic legacy.
PLATE 25. This satellite image of Earth from space shows the
major zones of dominant vegetation: forest, desert, mountains,
and other ecosystems. The continents here are seen as the raised
portions of enormous moving plates that have changed their
position over the history of the planet. Evolutionary biology and
the new plate tectonic-based geology complement each other
in broadening our view toward our living planet and its ancient
history. A unified biosphere, Earth's surface is the chemically ac-
tive aggregate of sunlight-transmuting, gas-exchanging, gene-
trading, environmentally transformative life forms.
 THE AMAZING ANIMALS

Can we believe that natural selection could produce,

on one hand, organs of trifling importance, such as the
tail of a giraffe, which serves as a fly-flapper, and, on the
other, organs of such wonderful structure, as the eye, of
which we hardly as yet fully understand the inimitable
perfection?

CHARLES DARWIN

Full fathom five thy father lies;

Of his bones are coral made:
Those are pearls that were his eye
Nothing of him that doth fade
But doth suffer a sea-change
lnto something rich and strange.
WILLIAM SHAKESPEARE

And, striving to be Man, the worm

Mounts through all the spires of form.
RALPH WALDO EMERSON

THE (BOWER) BIRDS AND THE (HONEY) BEES

Animals are multicellular assemblages that always develop from one

single cell. A swimming (undulipodiated) sperm penetrates and fer-
tilizes an egg.The fertfized egg then divides to form two, four, eight,
and more cells that make a blastula embryo, the essence of animal-
iry (fig. r3).
Each animal phylum (such as thejointed-foot Mandibulata, which

145
 sperm

."
J*f\-

sperm tail
(undulipodium)

FIcURE 13. Sexual lifehistoryof Dynasfes,abeetle.Phylum: Mandibulata.Kingdom:

Animalia. The hairy segmented structure is the larva, or grub, formed from the hollow
mass of cells known as the blastula stage of the animal embryo shown at right. The
blastula, a defining characteristic of all animals, develops from the sperm-fertilized egg,
which undergoes many cell divisions.
 The Amazing Animals 147

include insects, centipedes, and millipedes) is a great group-or a

formerly great group. The brachiopods, for example, are clamlike
but nonmolluscan creatures often found as fossils in Paleozoic ma-
rine rocks. Our own phylum, the Craniata, includes cartilagenous
and jawless fish as well as salamanders and pigeons. According to
one scheme, some thirty-eight phyla exist with animals that are alive
today; others have members that have long been extinct.
We have explored the origins of life, the bacterial realm, the pro-
toctists. Moreover, our discussion has been chronological, follow-
ing evolution and the rise in complexity. So, why now-well be-
fore the end of the book-do we talk of animals?
Fossils reveal that animals evolved earlier than plants or fungi.
Animals-exclusively marine animals-began leaving a rich fossil
record in the early Paleozoic. But there is no trace of plants or fungi
until more than roo million years after shelly animals appeared. Even
today, animals-typical in this regard, since life evolved in water-
abound more in water than on land. Only plants and fungi are pa:-
adigmatically creatures of the land. Microbes had to venture onto
land before these newer kingdoms could evolve there.
An animal cell is simple, relative to that of any plant. Plants, an-
imals, and fungi all sequester their genes into nuclei; all have mito-
chondria organelles to handle oxygen respiration. But plant cells have
the additional complexiry of an organelle for tapping the energy of
the sun. Animals lack such plastids. Nevertheless, members of King-
dom Animalia boast a proud heritage. Indeed, sensing and acting,
they have evolved some amazing capabilities.
Insectivorous bats detect their prey in the night skies by emitting
sounds at frequencies too high for humans to hear. They gauge, from
the echoes that come back to them, the presence and position of
objects in their surround. "Blind" when their ears are plugged, their
ultrasonic picture of the world may resemble our ultrasound scans
of unborn babies. Bat-blindness was discovered by the same soup-
boiling Lazzaro Spallanzani we encountered in chapter 3.
Not above incorporating colored poker chips into their con-
structions, the male bowerbirds of Australia and New Guinea build
 148 What ls Life?

elaborate colored mounds. Wings flapping and prancing, the bower-

bird calls loudly, flulling up his feathers, flar.rnting about. To assure
a female chooses him, he spruces up his ten-foot-square, foot-high
bower with bark, crushed fruit, charcoal, scavenged feathers, or even
blue laundry powder. To entice her, he decorates the bower with
bits of snail shell or fresh flowers he replaces daily. Nonetheless, when
she ventures in, he mates with her so violently it damages his own
bower, and then, when finished, he pecks, claws, and expels her from
his bachelor pad. This tragicomedy of lust is completed as the male
embarks on a proprietary strut about his territory, taking time out
only to destroy the bowers of younger males.
Some crabs acquire stinging sea anemones and wield them as
weapons against potential predators. Herring gulls drop clams,
whelks, and other shellfish from a height of several meters onto rock
to crack open their hard shells. Japanese green-backed herons lure
minnows by dropping twigs on pondwater. Dogs snifr, bark, and
scamper in their sleep as if chasing rabbits. Anesthetized and painted
with an obvious spot on their forehead while unconscious, chim-
panzees and orangutans (but, strangely, not gorillas) have awakened
to wipe their brow looking in a mirror-proof positive that they
recognize themselves. Subordinate simians hide their copulations
from dominant males. This suggests that they not only recognize
themselves but have a mental model of how others see them. Vervet
monkeys sound three distinct alarm calls, depending on whether the
object of their concern is a leopard, a python, or an eagle. Dolphins
whistle at an amplitude and frequency that is distinctive for each in-
dividual; they seem to call each other by name. Captive in an aquar-
ium, one bottle-nosed dolphin has been reported to irnitate, per-
haps for comic effect, the swimming postures of turtles and penguins.
This dolphin, who used a gull feather to scrape algae on an under-
water window apparently mimicked a human diver. The dolphin
even emitted a stream of bubbles like those the diver expelled.
But exceptional brain power is not always at the root of excep-
tional communicative and apparently intelligent animal behavior.
 The Amazing Animals I lnS

Honeybees see in color (including ultraviolet, which humans can't

see-but not red, which we can). They orient themselves by light
polarization in the sky. Cooling the hive by regurgitating water
droplets and fanning their wings, and warming it by shivering, they
are masters of environmental regulation. Honeybees seek pollen
grains and nectar and the workers that discover rich sources of these
materials communicate their finds to others. The successful scouts
appear to perform two distinct dances to specify the location: one
circular, for close food sources, the other a "waggle" dance, for food
more than about a hundred meters away. The assumption that this
instinctive bee behavior, the discovery of which earned Austrian-
German zoologist Karl von Frisch a Nobel Prize in r973, is no
more conscious than a computer program, has never been proven.
The honeybee's knowledge of dances seems to be innate, but it is
nevertheless possible that the bees are in fact "aware" of their
significance.
Animals are so amazing that we need not, as humans, consider
ourselves as anything more than animals in order to feel justifiably
proud. But as Donald Grifiin laments, ever since Darwin forced hu-
mankind to recognize its animal kinship, the inclination has been
just the opposite:

Much of twentieth-century science has gradually slipped into an

attitude that belittles nonhuman animals. Subtle but effective non-
verbal signals to this effect emanate from much of the scientific lit-
erature. Physical and chemical science is assumed to be more funda-
mental, more rigorous, and more significant than zoology. Modern
biology revels in being largely molecular, and this inevitabiliry di-
verts attention away from the investigation of animals for their own
sakes. Part of this trend may be due to an unrecognized reaction
against the deflation of human vaniry by the Darwinian revolution.
The acceptance of biological evolution and the genetic relationship
of our species to others was a shattering blow to the human ego,
from which we may not have fully recovered, for it is not easy to
give up a deep-seated faith that our kind is unique and qualitatively
superior.
 150 What ls Life?

Grifiin goes on to suggest:

a psychological palliative that may be subconsciously attracrive, even

to many scientists, is to shift attention away from the embarrassing
fact of our animal ancestry by accentuating those aspects of science
that are more akin to physics. This may help explain why so many
appear to be so certain that consciousness and language are uniquely
human capabilities and that the discovery of symbolic comrnunica-
tion by honeybees "upsets the very foundatron of behavior, and bi-
oiogy in general." Quite the conrrary, such discoveries in the field of
cognitive ethology extend and improve our understanding of animals;
a definition ofbiology that rules out rhose discoveries a priori suffers
from self-inflicted impoverishment. 1

Consciousness, private affair, is not directly measurable. But an in-

a

abiliry to render a qualiry measurable is no reason to assume its

absence-to assume that animals are mere instinct machines. Indeed.
we would take Grifiin a step further. Notjust animals are conscious,
but every organic being, every autopoietic cell is conscious. In the
simplest sense consciousness is an awareness of the outside world.
And this world need not be the world outside one's mammalian fur.
It may also be the world outside one's cell membrane. Certainly
some level of awareness, of responsiveness owing to that awareness,
is implied in all autopoietic systems. The world, afrer all, is nor a
petri dish; the sky does not rain agar. Every live being incessandy
senses and responds with alacriry to its surroundings.
A-11 animals have a multicellular, rnultirissued stage in their life

histories. But the complex chemistry of life is still confined to the

cell, minimally less than a single micron in diameter. Each gener-
ation of every kind of animal returns to the ancestral-type single
cell as fertilized egg. The increase in size and complexiry of ani-
mal life has occurred as socially aggregated cells evolved to form
unified bodies.
Animals eat other organisms. They do not enjoy the vegetal lux-
ury of feeding themselves through photosynthesis but must go out
into the world-begging, borrowing, and stealing-in order to ac-
complish what plants can do sitting calmly in one place. Many ex-
 The Amazing Animals 151

traordinary animal attributes stem from the requirement to procure

hard-to-get food. Others may be traced to the reproductive imper-
ative of sexuality: in each generation sperm and egg must join. The
aesthetics of bowerbirds, the dancing of honeybees, the dreaming
of dogs have all evolved. Connected to the procuring of food and/ or
mates, all such behaviors manifest the ancient autopoiesis of self-
perpetuating life.
Animal behaviors seem directed, "purposeful." Yet like bacterial
magnetotaxis and protist cannibalism, they may also be perceived as
ways in which the universe's dissipative systelru make waste of use-
ful energy by putting it in the service of localized pockets of stun-
ning order. Communication, imrtative learning, tool use, and con-
scious thought make sense from a thermodynamic perspective.

WHAT IS AN ANIMAL?

Most species of living beings today are probably animals. Estimates

range from about three to more than thirry million species in this
one kingdom. Some phyla, conspicuously successful, are well known:
the echinoderms (starfish, sea urchins, sea cucumbers), the mollusks
(clams, snails, squid), the craniates (fish, reptiles, birds), the coelen-
terates (hydras, corals, jellyfish), the mandibulates (insects and mil-
lipede$. Lesser-known phyla include the pogonophorans (some of
the deep worms), onychophorans (velvet worms on the South
sea
American forest floor), and pentastomes (which live in the nasal pas-
of mammals).
sageways
Animals-all with tlvo-parent sex and fertilized eggs that form
embryos-mature into reproductives subject to individual pro-
grammed death (see fig. t:). Despite their diversiry and exuber-
ance, animals are evolutionary newcomers. The first animals evolved
in an oxygen-rich world with large continental masses and open
seas-a world not too different from that which supports us today-
But at the time of their appearance 8o percent of the story of life-
thus far-had already been played out.
Ocean dwellers from the start, the earliest animals do not appear
 152 What ls Life?

in the fossil record until the end of the Proterozoic eon, some 60o
million years ago. The famous trilobites, early Cambrian marine an-
imals, are even more recent. Clear evidence of abundant animal fos-
sils with hard parts dates to fewer than 6oo million years ago. As
most animals do today, all inhabited seawater. Only a very few an-
cestors of modern animals-some craniates, worms, insects, spiders,
and mollusks-ever succeeded in leaving the ocean behind to sur-
vive on land (see plate r5).
Land animals-with their intricate bodies, devious minds, and
sometimes elaborate societies-seem to have evolved furthest from
the earliest cell. But consider: an animal is the very creature telting
this story of evolution. Might rhat teller be just a tad bit biased in
favor of its own kingdom? Perhaps, considering the source, the no-
tion of progress from "low" bacterium to "high" human is a delu-
sion of grandeur. As paleontologist Stephen Jay Gould once re-
marked, an intelligent octopus would probably regard eight arms as
more perfect than two.
The evolution of the first animals is a fascinating question. But
what is an animal? How might we have recognized the first mem-
ber of the new animal kingdom in its own day? Certainly an ani-
mal is not just an organism that moves but does not photosynthe-
size, because most bacteria and many protoctists would then, by
definition, be animals. What makes any animal distinctively and
uniquely an animal? What has a person in common with a worm,
a starfish, and a million kinds of beetle?
Animals, whether in the urban dark of a barroom or by a moon-
lit equatorial reef, all share the same life cycle. Fusion of the rwo
different-sized cells, egg and sperm, begins the process of animal-
iry. The sperm and egg merge into a single fertilized egg rhar divides
by nritosis to form the blastula. As cell division continues, the fer-
tile egg becomes the embryo-often a hollow ball of cells (see plate
16). Just as the embryo distinguishes the animal and plant king-
doms from the other three kingdoms, so the blastula embryo distin-
guishes the animal from the plant. A plant embryo is a solid mass
 The Amazing Animals 153

within the maternal tissue. A blastula is usually hollow; most animal

eggs develop into embryos in the water, free of maternal tissue.
Animal embryos assure the animal kind of individualiry. Even sex-
ual protoctists do not require sex to reproduce; moreover, they never
make the sexual product we call an embryo. Embryonic develop-
ment is not part of protoctist individualiry. Loose colonies of pro-
toctists tend not to have fixed body sizes and shapes; rather, pieces
of protoctists break off and form new amorphous wholes. Com-
posed of tissues with distinct connections between the cells-
desmosomes, gap junctions, septate junctions, and the like-the
bodies of animals are individualized. Such cell-to-cell connections,
unknown in any other kingdom, must be produced by embryo de-
velopment. In a predictable sequence, cells of the embryonic ani-
mal divide by mitosis, they roll over one another and set up alliances
such that many, sometimes most, of the animal's body cells must
die in a preprogrammed fashion. If the young embryo cells do not
die on cue or do not establish definite connections to send specific
signals through their junctions, no animal body develops. Animal
embryos are crucial.
ln animals the muscles, nerves, and circulating fluids such as blood
all derive by mitotic cell division and differentiation from the blas-
tula. Doomed to mortaliry, the blood, muscle, and nerve cells stop
dividing; their programmed death contributes to the construction
of the body and the continuing of the theoretically imrnortal egg
and sperm. Without the blastula embryo formed from the fertile
union of sperm and egg, animals would not exist. The blastula em-
bryo is the animal universal in all thirty-eight or so phyla. In the
usual course of animal development the cells of the blastula con-
tinue to divide, move, and die. They form the next stage, the gas-
trula, which makes a new mouth at the front end of a digestive tube.
The mouth is succeeded by a swelling, a stomach, and an anus in
this unfolding process, called, logically enough, "gastrulation."
The vast majoriry of blastulas in the thirty million animal species
"gastrulate" and end up with a distinctive "tube-within-a tube" di-
 154 What ls Life?

gestive system, committed to heterotrophic nutrition. Formed by

cell-to-cell communication via gap or septate junctions with nerve
cell synapses, the tube or gut establishes the ingestive nutrition so
nearly universal in the kingdom. But a few kinds of exceptional an-
imals lack intestines and even these must pass through a blastula stage.
This kind of embryo is the hallmark of any animal.
Neither fungi nor protoctisr, although mosr are heterotrophs,
forms any embryo at all. Probably correlated with the absence of
the embryo is the absence of animal-sryle distinctive individualiry.
Plant individuahry is far less fixed than that of animals. Although all
members of the plant kingdom do form embryos, plant embryos
are very diIl-erent from any blastula. Each plant cell is walled from
its neighbor, precluding the movements and realignments that all
blastulas undertake as they become individual animal larvae or adults.
An embryonic plant cell cannot form gap or seprate junctions, nerve
synapses, or any other of the animal cell-to-cell connections. Sta-
tionary in its place, the plant cell only grows, dividing by mitosis,
or dies. It is the fateful blastula rhat presages all rhe nuances of an-
imal behavior and distinguishes our kingdom from all the rest of
life.

GREAT-GRANDPARENT TRI CHO P LAX

To the bacterial realm goes the award for metabolic innovation. As

keepers of the biosphere, prokaryotes are the most inventive forms
of life, and their descendants include the now-essential organelles
within our own cells. The protoctists, too, dealt originally with the
problem of environmental threat: in autopoietically changing to
stay the same, they evolved new modes of programmed death, cell
fusion, sex. Many lineages invented metamorphosis: their watery
bodies developed-reversibly-into drought-resistant strucrures.
But with the origin of animals nature seems to have reached new
levels of playfulness, awareness, complexiry of form, responsiveness,
and deception.
 The Amazing Animals 155

A butterfly's wing bearing an imitation raindrop with a line dis-

placed just as if it had been refracted through real water; a cheetah
poised to pounce; an acrobat juggling upon a high wire: animals
anaze.
Today's minimal animal is Tiichoplax-a headless, tailless creature
discovered crawling on its belly along the side of a marine aquarium
in Philadelphia in 1965. Were it not for its sex life and its embryo,
Trichoplax would be a protoctist. Wafted along by its undulipodia,
Tiichoplax superficially resembles a slime mold slug or a giant ameba-
But it is a multicellular being throughout its life and is a true animal.
It has more undulipodia on its belly than its back. Having neither
head nor hind end, right nor left side, no eyes, no stomach, this min-
uscule slow crawler gives away the secret of its animality only at re-
production. After fusion with sperm, a spherical Tiichoplax egg be-
comes a blastula embryo that, with more cell division, flattens and,
ameba-like, slinks away. Although you probably wouldn't want to
hang a portrait of it in the drawing room, Tiichoplax likely bears a
strong resemblance to our earliest animal ancestors.
A sponge is an animal composed of only a few rypes of func-
tionally and morphologically different cells. For example, those on
the outside may grow glassy rods for support and protection; those
on the inside use their undulipodia to maintain a flow of water from
which food can be extracted. If yellow and orange (Haliciona)
sponges are squeezed through cheesecloth so that each is broken
into bits and mixed with the other, the cells manage to find their
own kind in the aqueous environment. After a few hours the cells
reorganize into fully formed and distinct yellow and orange sponges.
So too, a freshwater polyp, a jellyfish relative having about a hun-
dred thousand cells of a dozen cell rypes, can be disassociated into
single cells. In permissive solutions they begin to rearrange them-
selves.Unlike sponges, they cannot complete the process. Monstrous
growths, in which head, gut, and foot (basal stalk) realign unsuc-
cessfully, are the result. In this case, the integrating mechanisms as-
suring autopoietic self-maintenance fail.
 156 What ls Life?

In most colonial green algae and ciliates (all of which are pro-
toctists) any single cell may separate and reproduce on its own. In
others, only certain cells reproduce. The theme of animal evolu-
tion, the development of discrete individuals, involves curtailing re-
production in favor of specialization. Protoctist anarchies, in which
any cell could reproduce, were replaced during the emergence of
animals by cell oligarchies, in which only a few (sometimes a very
few) had the privilege of living on into the next generation by way
of progeny.
The transition from cell, to cell sociery to animal organism is an
old story in evolution: individuals group into societies, which
themselves become individuals. Under intense selection pressures,
swimrning protists became colonial protoctists. Then, in the later
Proterozoic eon, Tiichoplax-llke animal bodies appeared. The spe-
ciahzation of massive numbers of cells into integrated individuals is
at the base of animal life-and of those later groups, fungi and plants.

SEX AND DEATH

Only accidental, externally caused death exrsted at the origin of

life. So it was for a long time thereafter. But with protoctists came
"programmed death": death in which cells age and die as part of
the life of the individual. In farniliar animals-insects, mammals,
and birds-the difference between rhe part that dies and the part
that potentially lives on is the difrerence berween the body and the
sex cells. In mammals the sex cells (or "germ plasm," as biologists
sometimes say) are the only cells whose direct progeny survive into
the next generation. In contrast to the ova and sperm, 1fus ('sqrns"-
the animal body-has a discrete life span.
With a high degree of precision, animal cells must reproduce-
or cease reproducing. For example, during the intrauterine devel-
opment of the mamrnal brain more than 90 percenr of the cells that
develop die before the fetus becomes an infant. These brain cells
stop growing and disintegrate, are sacrificed in the process of grow-
 The Amazing Animals 157

ing a healthy infant. The essential difference berween the living germ
cells and the dying body cells of animals is likely very old.
We speculate that the ancestors of animals were composed of rel-
atively few cells that differentiated into at least rwo distinct kinds.
One kind specialized in using their 9(z)+z microtubule organelles
to form undulipodia for propulsion, for sensing prey, for fostering
water flow over or through the animal, or for sweeping food par-
ticles into and along digestive systems (see plate I7). But it is an
oddity of physiology that once animal cells dedicate their centrioles
to forming undulipodial shafts, they can no longer use them to cre-
ate the motility apparatus for mitotic cell division. This means that
animal cells stood to gain by sticking together in specialized colonies.
Even today an animal cell, whether of a grow-
tissue or a sperm after
ing the undulipodium, no longer divides. A centriole forms a kinet-
osome and relinquishes cell immortaliry; a kinetosome cannot revert
to a centriole. The irreversibiliry of kinetosome formation apPears
to be an inviolable rule within the animal kingdom. Animal cells
can either form kinetosomes (grow undulipodia from centrioles) or
reproduce by mitosis-but not both. An anim.al cell with a kineto-
some is a dead animal cell-its days are numbered, as it will not di-
vide again.
Perhaps the DNA reported by David Luck and John Hall to be
in the kinetosome-centriole is used for mitosis or to form an un-
dulipodium but not for both. Like choking from inhaling water, any
attempts of cells to simultaneously reproduce and maintain un-
dulipodia would have been thwarted. And yet animals seem to have
found an answer to this genetic dilemma: by sticking together in
colonies-colonies where some cells reproduce while others form
undulipodia-they could in effect have their cake and eat it too.
The restriction of a cell unable to divide after growth of its 9(z)+z
organelle was overcome by colony formation. The great majoriry
of cells retained their option to divide, while a few sacrificed im-
mortaliry to be undulipodiated. But even the cells that do divide in
the animal do not do so indefinitely. After 6oo million years the
 158 What ls Life?

adult animal is still a mated protist's way of making other mating

protists.
Our whole life from womb to tomb is in fact an interim stage in
the life cycle of tiny fused cells. Animals emerge into another di-
mension, visible life and consciousness, only to return via sex to their
ancient single-celled, microbial state. Death is the price we all pay
for this ancient history of multicellular compounding, for this in-
ability of hungry protists to undo their Proterozoic entanglements.
-What
"dies" is the body, the adult flesh after it has released into the
water or body fluid the protist-like tailed sperm and chubbier egg.
Animal life did not appear de novo, but from protoctist predeces-
sors. Protoctists with elaborate cycles of fertlTtzation, multicellular-
iry and meiosis became animals.
Like programmed death, gender is not intrinsic to life. Gender
evolved. Cells of different mating rypes, like protocrisr lovers today,
were initially identical to each other in appearance. The seasonal
merging and restoration of chromosomal numbers in Grtilization
set the stage for the origin of gender. The first mates met slapdash
in a watery environment then, as protoctists do today. Responding
to slight chemical differences in each other, mates came together.
Sponge, sea urchin, fish, and even mammalian sex cells, like their
protoctist ancestors, still meet in watery places.
Animal cells continue their ancient practice of aquatic encounter.
The sex cells of oysters and even some frogs and fish meet directly
in the water, to fuse unattended by adult bodies. In reptiles, birds,
and mammals, however, sexual mergers occor in viuo. Genttals
evolved independently in many animal lineages. The penis or in-
trornittent organ of the male created a delivery system for sperm.
The female genital tract afforded the ova a protected place where
fusion could occur. The many small sperm of males compared to
the few larger eggs of females was the beginning of an evolution-
ary asymmetry which today expands into the realms of political, so-
ciolinguistic, and psychological debate. Evolutionary biologists sug-
gest that early sexual inequality-males maximize reproduction by
 The Amazing Animals 159

inserninating the largest possible number of females, whereas after

a certain lirnit mating becomes superfluous to females constrained
by devotion to their lesser quantiry of eggs-is behind distinct male
and female attitudes toward sex.

CAMBRIAN CHAUVINISM

English geologist Adam Sedgwick (r785-r 873) named the time pe-
riod to which the oldest fossils belonged Cambrian, after "Cam-
bria," the old name for Wales in southwestern Great Britain. To him
and other early paleontologists the appearance upon Earth of ani-
mals seemed miraculously sudden. A11 prehistory prior to Sedgwick's
Cambrian became known as the "Precambrian." I-Jntil the late twen-
tieth century the origin of the Cambrian animal fossils was consid-
ered "the most vexing riddle in paleontology."2 So quick was the
apparent appearance of animal life in the fossil record-not only in
Wales but also in New{oundland, Siberia, China, and the Grand
Canyon of Arizona-that it is still referred to as the "Cambrian ex-
plosion."
Today, much of the answer to the riddle is known. Soft-bodied
and other inconspicuous protoclists, so-called "protozoa" whose fos-
sils were once dismissed as those of tiny invertebrate animals, in fact
preceded animals by at least 5oo million years. Because, Iike early
animals, most protoctists were small and did not form hard parts,
they remained largely undetected and unpreserved. Life prior to the
Cambrian, despite its astounding biochemical and metabolic inno-
vations, is still often dismissed as "Precambrian"-with the conno-
tation that nothing much worth mentioning in evolution happened
bervrzeen the origin of life and the appearance of shelled animals.
Bacteria and protoctists set the stage. They, not animals, introduced
DNA recombination, locomotion, reproduction leading to expo-
nential growth, photosynthesis, boil-proof spores. They, not animals,
pioneered symbiosis and the organization of individuals from mul-
ticellular collectives. They invented intracellular motiliry (including
 't60 What ls Life?

mitosis), complex developmental cycles, meiosis, sexual fusion, in-

dividualiry and programmed death. The prokaryotic microbes, not
animals or plants, still run all the geochemical cycles that make the
planet habitable. The protoctists, in their new status as individuals
from coevolved bacterial communities, invented resistant cysts,
skeletons and shells, gender behaviors, cell-to-cell communication,
lethal toxins, and many other processes later co-opted by animals.
Animals were preceded by bacteria and protoctists, not by chemi-
cals. The animal explosion had a long microbial fuse.
Patterned fossils of tiny shelly plates, known collectively as scle-
rites, mark the beginning of the Cambrian about j4r million years
ago. The lowermost time-rock division or period of the Phanero-
zoic eon is the Cambrian. Distinctive animal fossils abound in Pha-
nerozoic rocks overlaying barren Proterozoic strata. Sedimentary
rocks from all over the world deposited 5 ro million years ago (still
in the Cambrian) contain a striking array of skeletalized marine an-
imals. At about this time, brachiopods (lampshells) and annelid
worms appeared. So did trilobites and other joint-footed animals
(of which insects and lobsters are modern examples).
Some paleontologists still wonder how these various phyla could
have cropped up "all of the sudden." Noting iron rust and other
clues that oxygen had entered the atmosphere 2,ooo million years
ago, some scientists suggest that a threshold level of atmospheric oxy-
gen (Oz) itself induced animal evolution. But any scenario to ac-
count for the "sudden" appearance of animals is almost surely a nris-
reading of the evidence. Animals, although they evolved late in the
history of life, did not evolve sr"rddenly. "Seemingly," writes pale-
ontologist Harry B. Whittington, "there was a long period of meta-
zoan [animal] evolution before the Cambrian, but it is only in the
eariiest Cambrian rocks that minute shells of metazoans appear. . . .

The Burgess Shale shows that it was not only metazoans with hard
parts that were diversifiiing in the Cambrian, but also soft-bodied
metazoans, including coelenterates, worms, arthropods, chordates,
and various strange animals."3
 The Amazing Animals 't 61

The Burgess shale is a collection of Cambrian fossils exposed on

a high mountain in Yoho National Park in British Columbia,
Canada. Discovered by Charles Walcott in r9o9, the exquisite and
numerous Burgess shale fossils have given paleontologists lifetimes
of work. Because even soft-bodied animals were preserved in it, this
shale is a treasure. Shallow marine dwellers were preserved in the
underwater mudslides that made the Burgess shale. These were a large
variety of organisms, some similar to modern forms, others with
no known descendants. Among the beautiful, if monstrous, animals
are Opabinia, a five-eyed sea-bottom crawler with curved tail fins
and a grasping, jointed organ that suggests it was a formidable pred-
ator, though only four inches long. Hallucinogenia, in accordance with
its name, has puzzled paleontologists, because until recently no one
was sure which side was up (spikes as armor) and which side was
down (spikes as legs). Of the many Cambrian arthropods that the
Burgess shale has preserved, only one sort gave rise to a lineage that
much later evolved into the vast array of land creatures with six legs
known today as insects. Had evolution taken another course, an-
other Cambrian arthropod-or an entirely different animal, for that
matter-might have gone on to populate the continents.
Among the most touching of the Burgess specimens ts Pikaia, the
first known member of our own chordate group-the one to which
hunrans and all other animals with backbones belong. Pikaia, a seg-
mented wormlike swimming creature, is inconspicuous compared
to more spectacular Burgess forms. But it had a solid cartilaginous
rod-the notochord-running down its back. This universal struc-
ture of chordates if not present in the adult is fleetingly apparent in
larvae or other immature life cycle stages. Until the discovery of
Pikaiarnthe Burgess shale, no chordates were known from any rock
older than about 45o million years, deposited during the Ordovi-
cian, the geological period after the Cambrian.
The Burgess chordate is a stunning discovery, because it shows
that the predecessors of darting Pikaia-which may have been the
ultimate ancestor of all fish, amphibians, reptiles, birds, mammals,
 162 What ls Life?

and us-were alive and navigating the muddy warers 5ro million
years ago. The success of Pikaia may be directly responsible for the
later emergence of such a wide diversity of forms, from mugwump
to turtle, moose, rabbit, and giraffe. The presence of streamlined
Pikaia, with its poinry somewhat cobralike flattened head and bi-
furcating snail-shaped tail, shows that our ancestors swam in the
primeval oceans.
Before the armor-headed trilobites crawled the planet, before
droves of clamlike lampshells expired in Cambrian muds, before eu-
rypterid "sea scorpions" left their hard exoskeletons in the fossil
record, soft-bodied animals proliferated. Even less obvious and far
older than the Burgess shale animals are the "Ediacaran" beings pre-
served in sandstones 7oo million years old-before the Cambrian
period, before the Phanerozoic eon. Most are probably not animals
at all but bizarre, extinct protoctists. In the r95os Martin Glaessner
of the lJniversiry of Adelaide named these extraordinary fossils af-
ter a rock formation in the Ediacara Hills of Australia. Similar soft-
bodied beings have been found inEngland, Namibia, Greenland, the
coast of the White Sea in Russia, and some rwenty other localities.
The Ediacaran organisms seem to have been floating gelatinous
beings enjoying shallow water at sandy beaches. Some were flat, oth-
ers "quilted," others intricately textured organisms. They ranged,
shapewise, from leaflike Pteridinium to rhree-armed Tribrachidium.
But these Ediacaran beings seem to have formed no hard parts, eggs,
sperm, or blastula embryos. They may have been large protoctists,
animals, or both. Some of the larger Ediacaran beings probably pho-
tosynthesized in shallow coastal seas. Others fed on bacterial pas-
tures. But their lack of armor indicates that large predatory organ-
isms had not yet evolved-it was truly a "Garden of Ediacara."a
Ediacarans may have been ancestors to the Burgess Cambrian an-
imals or, more likely, because they are utterly unique, they may have
been one of evolution's many "false starts." The earliest marine
animals-whatever these were-may have fed on protoctists, in-
cluding algae. The small size and relarive mobiliry of these algae
 The Amazing Animals 163

eaters probably guaranteed a nutritional niche without much com-

petition. Only after animals began trying to eat one another-and
evolving larger size and hard bodies as defenses-did they become
obvious in the fossil record. Mating, embryo-forming animals must
have exploited resources for millions of years before they evolved
hard, easily preserved parts. Cambrian fossils are only the tip of the
iceberg of animal evolution. They were hardly "the first animals."
A thermodynamic truth is that as heat dissipates life organizes and
its surroundings degrade. There is no life without waste, exudate,
pollution. In the prodigaliry of its spreading, life inevitably threat-
ens itself with potentially fatal messes that prompt further evolu-
tion. But sometimes waste can be fashioned into something useful.
Within protoctist or animal cells the concentration of calcium
ions-Ca++ charged particles-is some ten thousand times less than
that of seawater. Calcium phosphate becomes a fatal precipitate as
"rocks" form inside cells when too much calcium enters. And by
binding with phosphorus (as phosphate), errant atoms of calcium
deprive the cell of an essential ingredient for making DNA, RNA,
and membranes. By contrast, in controlled, small quantities calcium
ions can be an intracellular resource. Poison depends on dose. In
small doses calcium ions signal; indeed, they are part of the elec-
trochemistry of thought. But the calcium excess must always be
shunted outside of the cell. If the process of extruding calcium be-
yond cell frontiers lapses, then chemistry takes over. As people with
kidney stones know only too well, calcium phosphate, where it
doesn't belong, causes trouble. While animals remained soft they re-
leased calcium into seawater. Near the beginning of the Cambrian
period in the Phanerozoic eon, however, some began to control their
calcium extrusion.
Evolving, early animals converted potentially threatening block-
ages into living architecture. Our bones and skulls, like those of the
fish-headed amphibians that preceded us, are still made of calcium
phosphate salts (CazPO+). Some animals, such as corals, fashioned
calcium, phosphates, and carbonates (CaCO:) as their exteriors.
 164 What ls Life?

Other organisms deposited calcium inside as teeth. Replacing or-

ganic cartilage, hard calcium phosphate infiltrated the proteins to
provide the structural framework (shells and bones) attached to mus-
cles. As armor emerged in some Cantbrian creatures, teeth and pok-
ing appendages evolved in others to penetrate that armor.
Human industry has no monopoly on hazardous waste. Earlier
liG forms tell us by their examples that long-term survival involves
not so much halting pollution as transforming pollutants. Ternrites
build nests from feces and saliva. Pollution in the form of calcium
excrement, patched and reworked by the busy muscles of animals,
formed the basis for the first shells.

EVOLUTIONARY EXUBERANCE

Becoming numerous, using their powers of movement to colonize

new territories, animals evolved, thus creating and inhabiting new
ecological niches. The Russian-American novelist and respected en-
tonrologist Vladinrir Nabokov (r 899-r 977) once suggested that pat-
terns of certain butterflies look more like the whimsical touches of
an artist god than traits that had merely blindly evolved. Yet with a
fuller, less mechanical view of evolution such animal traits can be
explained.
Evolution is no mechanical law but a cornplex of processes, sen-
sitive and synrbiogenetic, in part resulting from the choices and
actions of evolving organic beings themselves. Natural selection
is often said to "fayor" this or that trait. But the nature that se-
lects is largely alive. Narure is no black box but a kind of sentient
symphony.
Of all the organisnes conceived, spawned, hatched, and born, very
few survive. tstes in food and mates lead certain beings ro gener-
ate more offspring than their fellows produce. In some cases females
help designate the genetic makeup of populations by choosing the
healthiest, most ostentatious, or strongest males. But conscious in-
tervention in evolution by those so evolving can be of a more sub-
tle variery. A butterfly's wing with an imitation raindrop showing
 The Amazing Animals 165

a line displaced as if it had been refracted through water does not

require a conscious author, and yet it could arise from conscious-
ness. Beguiling5 artifices might arise from misperception by intelli-
gent actors, for example, birds consistently mistaking an insect's wing
pattern for a leaf. Nature is made partly in the image of mind.
Nabokov was right to say that the greatest enchantments in both
art and nature involve deception. The element of surprise is the
revelation that a given phenomenon of the environment was, un-
til this moment, misinterpreted. Animals who experience surprise
as a pleasure are likely to recognize carnouflage and leave more
ollspring than are their less perspicacious brethren. Selection as
nature, filled with live, sensitive beings, is by no means blind (see
plates r8a and I8b).
Deception is very important in animal societies-so much so that
some sociologists speculate that human technological intelligence
is an evolutionary offshoot of "Machiavellian" social intelligence-
the capacity to procure food, mates, child care, and so on by out-
witting others in the tribe.The ourwitting, outrunning, or outfight-
ing need not be entirely conscious. Fearful apes and monkeys
undergo a physiological reaction that stands their hairs on End. The
effect to a potential combatant is an increase in size of the oppo-
nent, commanding if not fear then respect.B-52, punk, and other
"big hair" hairsryles may similarly affect onlookers.
Since as naked primates we humans reveal more skin than hair,
our own hair-raising response is rather measly: a prickly feeling, a
hot flush across the nape of the neck, a tingling along the spine.
Gooseflesh seems to be the evolutionary vestige of follicles that still
go through the motions. Gooseflesh nevertheless exemplifies the
evolutionary link between body and mind. A goosebump-mediated
increase in a mammal's apparent size would be useless in a senseless
world. But we live in a sensuous one, where details determine food
and mate choices that in some cases spell the difference berween life
and death, between procreation and barrenness.
One of the sublime mysteries of life-supposedly calling evolu-
tion into question by its very presence-is the eye. Darwin wrote
 '166 | wf,ut ts Life?

of the eye's "illimitable perGction." The eye, connected to the brain,

perhaps seems perfect because it is the evolutionist's principal tool.
But how rnight the eye, this subtle source of perspectival and reflex-
ive mystery, have evolved?
The problem at first sight seems impossibly difficult. But not if
we remember the microbes. Vision was anticipated in light-sensi-
tive bacteria. Rhodopsin, "visual purple" of the mammalian retina,
is a colored protein complex present in abundance in the pink, salt-
loving archaebacteriurn Halobacter, where it is equally sensitive to
light. The colored pigment portion of rhodopsin is retinal, a chem-
ical similar to carotene of carrots and formed by the oxidation of
vitamin A. Retinal, the absorber of light in the retina of the mam-
malian eye, has a 3,5oo-million-year history.
Using a plastid inherited from a cyanobacrerial ancestor, the di-
nomastigote Erythrodiniun functions as a kind of single-cell eye.
With its "imitation lens" and "imitation retina," this protist evolved
a light-sensitive focusing device that involves most of its tiny body.
Insects, flatworms, sea slugs, and frogs have eyes that are very differ-
ent from one another, but all have carotene-derived light-sensi-
tive membranes, lenses, and movable parts that direct light signals
to locomotory organelles (such as undulipodia) or tissues (such as
nruscles). Some evolutionists suspect eyes evolved in more than
forty distinct lineages of animals. In all, light sensing is connecred
to movement in some way so that, once signaled, the creature can
respond.
Sight, and its organ, the eye, may seem miraculous. Nonetheless,
eyes exist along a continuum of complexity that both precedes mam-
mals and is lately surpassing them, in forms such as infrared detec-
tors, radio telescopes, and satellite imaging technology. Light sensi-
tiviry in the rudimentary sense, even antedates life itself colored
compounds react in higtrly specific ways to visible solar radiation.
The human eye bears the marks of its microbial predecessors in
other ways. The "rod" and "cone" cells are incapable of mitotic di-
vision; they bear kinetosomes and short undulipodia, inherited from
their protist ancestors. The bony socket in which the eye resides de-
 The Amazing Animals I lel

rives from the autopoietic necessiry of recycling calcium waste. Over

time, life becomes more organized, integrating chemicals and even
waste into beings so sensitive they eventually begin to perceive their
own condition.

MESSEN GERS

By the late Devonian the phyla Mandibulata, Annelida, and Crani-

ata had evolved representatives able to survive the rigors of life on
land. Living beings evolved in water. The bacteria and protoctista
cells since their inception were bathed in fresh and salty water. Des-
iccation was a dire threat any land pioneer had to overcome. Evo-
lution of terrestrial species was no meager accomplishment. But the
evolution of land animals was a triumph not just for individual or-
ganisms and species; it was a victory for the biosphere.
Movement and intelligence permitted land animals to act as vec-
tors and messengers, to spread themselves to once-remote regions.
By the earlyTertiary birds had begun distributing phosphorus, a lim-
ited resource, to northern lakes and alpine peaks-simply by eat-
ing in one area and excreting in another. Carrying archaebacteria,
ciliates, and other microbes in their rumens, cowlike animals digested
grass, releasing methane, a greenhouse gas, into the atmosphere. Ni-
trogen-rich animal excrement accelerated algae growth, and fed fish
and copepods in cold-water ecosystems. Especially during the
Cenozoic, the most recent 65 million years, the quick reaction times,
continent-crossing migrations, and complex social interactions of
animals have accelerated activities within the biosphere.
But long before the Cenozoic, life was also a geological force. Pho-
tosynthesizing blue-green bacteria retained water in soil and sand,
makingEarth's surface green with chlorophyll. Carbon-precipitating
life sequestered more and more carbon into coal and limestones, con-
verting a tepid planet into one that indulged in glacial episodes. Land
life created soil from planetary rubble. Ocean life transformed salts
into reefs and evaporite flats.
"Proprioception" is the word for an animal's sensing various parts
 16A What ls Lif e?

of its own body. Human beings today, likely the most populous of
all mammalian species and certainly the most widespread, together
behave as a kind of planetary proprioceptor, giving the biosphere
sensations of itself. The greatest diversiry of life exists in tropical
jungles, such as the Amazon rain forest. Considering that distinct

rypes of bacteria have merged to form the eukaryotic cell, and that
colonies of eukaryotic cells evolved into animals, one wonders what
may result from interactions in dense, animal-rich conrrnunities.
Just as animal flesh was honed from the raw material of bacteria
over eons, so complex interactions produced fledgling individuals
at a scale beyond that of animals. Ants, termites, and bees form so-
cieties that carry out works in common. Reminiscent of human civ-
ilization, these insect workers methodically care for the young and
divide labor among specialized castes of soldiers, workers, and re-
productives. But whereas human civilization is only several thou-
sand years old, fossil evidence shows that ants and bees have been
organized into collectives for at least 4o mrllion, and termites for
perhaps zoo million years.
Together, animals confer their powers of movement and percep-
tion on the biosphere, making it an orgarized collective, the largest
organic being of all. The animal acrors of the global hive are at least
6oo million years old. Snakes sense infrared radiation. Whales hear
ultrasound. Bees detect the plane of polarization of visible light.'Wasps
see ultraviolet light patterns in flowers that look unpatterned to us.
Dogs enjoy "ultrasrnell." Sharks Grrer our buried prey by detecting
electrical potentials from the heartbears of the hidden. Animals sig-
nal, and sense, and engage each other and their living environment
in the visible, auditory, olfactory, and invisible radiative realms. Such
sensitiviry so widely dispersed, sensitizes the entire biosphere.
Humans have extended one version of animal sensitiviry into near
Earth orbit. The image of Earth from space expands our awareness
of the global environment. From the rudiments of animal sensibility
and movement have come technological instrumentation, wheeled
vehicles, and telecommunication. Together, the eyes of blackbirds,
 The Amazing Animals 169

FIGURE 14. Eschiniscus blumi, a "water bear." Phylum: Tardigrada. Kingdom: Ani-
malia. These microscopic animals, named water bears by English naturalist Thomas
Huxley, are known as tardigrades. Highly sensitive to their boggy environments, they
survive drying out in temperature ranges from 150oC to -27O"C. These microbeasts
occur all over the world, but because the largest are no more than 1.2 mm in length,
they remain obscure. The span from claw to claw in the photo is fewerthan O.5 mm.

the sonar oi bats, the heat absorption of rvorms, the bacteria-derived

lur.ninescence of nrarine fish, and the aggregate awareness of un-
tr>ld nunrbe rs of rvalkinq, crawling, flying, burrorvins, thinkine be-
ings produce lnore than the sum of the parts. Sensitivities interact.
There are responses to resporlses. Animal awareness is not only a
straightfbnv:rrcl accumulation of eyes, ears, touch, and other senses
but an incalcul:rble synesthesia of nixed senses whose wholeness can
be but gleanccl by the hunran conscioustress, rvhich forms only :r
part (fig. I,+).
 17O I wr,ut ls Life?

Both the French paleontologist-priest PierreTeilhard de Chardin

and the Russian atheist Vladimir Vernadsky agreed that Earth is de-
veloping a global rnind. This layer of thought in the shape of a sphere
they called the noosphere, from Greek noos, mind. The aggregate
net of throbbing life, from flashing fireflies to human e-mail, is the
developing planetary mind. Perhaps, like the brain of a human babe
with many synaptic connections that diminish over time, the noos-
phere is still in its infancy. Polymorphous, paranoiac, confused, yet
intensely imaginative, the thinking layer of Earth that is largely the
unexpected product of animal consciousness, may now be in its most
impressionable stage.

SO, WHAT lS LIFE? Life is evolutionary exuberance;it is what hap-

pens when expanding populations of sensing, active organisms
knock up against each other and work things out. Life is animals at
play. It is a marvel of inventions for cooling and warming, collect-
ing and dispersing, eating and evading, wooing and deceiving. Life
is awareness and responsiveness; it is consciousness and even self-
consciousness. Life, historical contingency and wily curiosiry is the
flapping fin and soaring wing of animal ingenuiry the avant-garde
of the connected biosphere epitomized by members of Kingdom
Animalia.
 FLESH OF THE EARTH

I hold that the fruit of the Tree of the Knowledge of Good and
Evil was Soma, was the kakuli, was Amanita muscaria, was the
Nameless Mushroom of the English-speaking people. The Tree
was probably a conifer, in Mesopotamia. The serpent, being
underground, was the faithful attendant on the fruit.
R. CORDON WASSON

Truffles . . . are to be called certainly nothing other than flesh

of the earth. Best at spring and more often during thunder,
they are said to arouse dying love.
FRANCISCUS MARIUS CRAPALDUS

THE UNDERWORLD

Academics still often partition life into zoology, the study of ani-
mals, and botany, the study of plants. But what of pink molds, single-
celled yeasts, puffballs, morels, and psychedelic mushrooms?
Fungi have been lumped with plants because they aren't animals.
Medieval scholars working in a three-kingdom system suggested
they were zombielike, half-dead forms straddling the mineral and
plant kingdoms. lJntil quite recently the scientific term for fungi
has been Mycophyta-from Greek mykes (fungus, akin to mucus)
andphyton (plant). Although none photosynthesize, like plants, some
fungi are rooted. But they are best classified as unique in their own,
solely fungal kingdom: the Mychota. 'And fungi were fungi," wrote
theJapanese poetJunThkami, "they're like nobody else onEarth."l
In the English-speaking world the protorypical fungus is a dark,

171
 172 What ls Life?

dank toadstool, dimly associated with witches, smelly feet, and re-
frigerators, and generally to be avoided. "Fungi," declared the eigh-
teenth-century French botanist S. Veillard, "are a cursed tribe, an
invention of the devil, devised by him to disturb the rest of narure
created by God."2
Fungi do require sex to form the morel or the mushroom, but
they can reproduce without it. Because they don't photosynthesize,
they can live in utter darkness. Their vampiristic existence often re-
quires them to do so-sometimes on rather scarce resources of food
and water. Reversing the animal technique of taking in food and
then digesting it, fungi digest food outside their bodies. They then
absorb the nutritious particles through their membranes.
Fungi differ from all other life (fig ,S).Unlike plants and ani-
mals they form no embryos. They grow from tiny propagules, pack-
ages called spores. lJpon moistening, the spores form threads, thin
tubes, the hyphae. Yeast cells (used in brewing beer and raising bread)
bud offsingle cells. Lacking the whiplike structures of undulipodia,
neither single-celled nor multicellular fungi ever swim. Some, called
by the fancy name laboulbenomycetes, indulge in fungal sex to form
spores that disperse on insect legs. Spores of others attach to ntam-
mal fur, are sneezed out, or drift in wind. When hitchhiking spores
come to rest and sense moisture, hyphae begin to grow up, down,
and sideways. Like plants and animals, fungi are made of nucleated
not animals) they possess tough cell walls. Fun-
cells. Like plants (but
gus cell walls are made of chitin, a nitrogen-rich carbohydrate; plant
cell walls are made of cellulose. Many fungi have passages in their
cell walls that allow mitochondria, nuclei, and other organelles to
move between cells. Some lack cross walls altogether and are more
a growing mass of tubes than multicellular individuals.
Fungi break down dead and sometimes live bodies. For more than
4oo million years they have been settling and growing on a huge
variery of foodstufls other organisms eschew. A few grow in the sea
or underwater, but they are basically landlubbers. Fungi were
among the first organisms to make use of terrestrial environments,
basidiospore
P
-an
development \9,
E

lclamp"
connection,
indicating
fungal sex
has occurred

FIGURE 15. Different stages in the sexual life history of an Amanita fungus. Coun-
terclockwise from bottom: mushroom; close-up of sexual tissue or basidia found in
the gills; basidia giving rise to drop-like basidiospores; hyphae that grow from spores;
nuclei passing through the hyphae in the fungal sex act known as conjugation.
 174 What ls Life?

enabling the development of many other land dwellers. That fungi

prefer land was proved when scientists returned to Aluin, a sphere-
shaped submersible vehicle whose mission to the bottom of the
ocean ended when its lifeline to the mother ship snapped. Two years
later an intact, if soggy, sandwich was discovered on board the sunken
scientific vessel. In contrast, even inside a lunch box any sandwich
would be devoured if it were abandoned for rwo years on a picnic
table at Earth's surface. Fungal spores are found virtually every,where
in the air but do not grow on the sea bottom, perhaps owing to an
overabundance of salrwater.
Converting waste and corpses into resources, making nutrients
available to land life, fungi are invaluable to global metabolism. Fungi
thrive amid nature's informal cofiins as undifGrentiated masses; each
fungal thread is bounded, but the expanding organism as a whole-
a system of tubes-has no clear borders. Fungi, with their threads
penetrating food today but excised by environmental contingency
tomorrow, are truly fractal organisms.
The fungal Gature commonly called toadstool is really only the
rninuscule tip of an amorphous underground web of living threads
("hyphae"). A mold growing on bread or fruit demonstrates the typ-
ical fungus's lack of regular boundaries. The fungal thallus, an old
word for plant tissue undifferentiated into root, stem, or leaf, is also
called a rnycelium or mycelial ner\,vork. As the verb "dial" has come
to mean "depress buttons on a touch-tone phone," so botanical
terms-fruiting body, spore-applied to fungi persist despite their
inappropriateness.
Lacking discrete borders, fungi sprawl. Consider the tree-root col-
onizer Armillaria bulbosa. This organic being locates fresh food by
continuous exploratory growth. A fungal clone, able to recognize
itself from other expanding subsurface fungi, Armillaria has terri-
torialized the forest underworld with such inexorable fortitude that,
after an estimated r, joo years of growth, one sample growing be-
neath virgin coniferous forest in Crystal Falls, Michigan, now en-
compasses some thirty-seven acres. This individual fungus has an es-
 Flesh of the Earth '17 5

timated weight of more than eleven tons. A single individual, its

genes have been sampled and compared throughout and determined
to be the same. Such genetic stability is impressive. Fire, forest suc-
cession, and changes in food availabiliry have isolated parts of the
sprawling Armillaria, which, nonetheless, maintains its genetic in-
tegriry. Are the broken-offpieces separate organisms? Or should they
be regarded as the dispersed limbs of a single subterranean being?
Reminiscent of a Stephen King novel, the great chunks of biomass
continue to thrive, unperturbed by their multiple amputations.
"Although fungi have acquired notoriery as agents of disease and
as producers of psychedelic toadstools," writes scientist Clive Brasier,
"their vegetative structure, the mycelium or fungal thallus, has a
somewhat lower public profile."3 The low profile is literal, as most
of the mycelium exists in an extensive network, out of view be-
neath the soil. Great mycelia of foraging hyphae thrive beneath the
forest trees. The living threads called hyphae tend to fuse. After "hav-
ing sex," they eventually form mushrooms or mold tissues that in
turn undergo meiosis, forming spores. These disperse over forest and
field to grow again in search of mates.
Each mycelial nenvork is a fungal clone, the far-reaching offspring
of a single genetic line. Above ground, fungi produce airborne
spores, some of which you no doubt are inhaling now. When they
land, the spores grow wherever they can. Sprouting netr,vorks of
tubes, hyphae, into moist substratum, the fungi once again produce
copious quantities of disseminating spores, spreading their strange
flesh through the soil they help create.

KISSING MOLDS AND DESTROYING ANGELS

No one knows how many species of fungi exist. Some say a hun-
dred thousand. Others estimate one and a half million. Mycologist
Bryce Kendrick of 'Waterloo (Jniversity in Canada claims that to-
day fungi are more diverse than plants but less so than animals.
As with the other four kingdoms-Monera (bacteria), Protoctista,
 176 What ls Life?

Animalia, and Plantae-Mychota (fungi) have been arranged into

phyla. There are five major groups or phyla of fungi. The zygonry-
cotes (from Greek zygon, "trvin" or "pair") or mating molds lack
cross walls separating their cells. Mitochondria and nuclei travel eas-
ily through their open hyphal tubes. In zygomycote mating, special
hyphae-the gametangia-grow toward one another and fuse.
From this fusing of gametangia comes the production of resistant
spores. Once the hyphal ends lock, nuclei flow through the tubes
andjoin, probably in pairs. When the mating is over, meiosis oc-
curs, producing darkening spores in the head of a black spore holder.
Rhizopus stolonifer, the most common of the black bread molds, is
one example of a zygofungus.
Most molds (such as the pink bread mold l,leurospora, Clauiceps,
and ergot fungi) and most yeasts (such as Rhodotorula and Saccha-
romyrcs) are ascomycotes. They form asci, sacs or capsules that de-
velop when hyphae of compatible genders "kiss" and permanently
fuse. The complex tissues and sexual spores produced from such li-
aisons is "mold." The hyphal threads, the nonsexual fungal body
parts, are there but invisible to the naked human eye.
Ascofungi eat and degrade resistant plant and animal compounds,
such as the cellulose and lignin of wood, the keratin of fingernails,
and the collagen in mammal bones and connective tissue. By break-
ing down such compounds, these fungi release carbon dioxide, am-
monia, nitrogen, and phosphorus to the rest of the biosphere. The
evolution of wood placed great selection pressure on land-dwelling
fungi to invent ways to degrade lignin, thus ensuring a coevolved
biospheric cycling of matter. Some scientists posit that a lag in fun-
gal evolution contributed to the worldwide accumulation of coal
in the late Paleozoic era.
Humans ever).where are most farniliar with members of the third
fungal phylum (see plate r9a). The basidiomycotes have reproduc-
tive structures called basidia, which resemble clubs (basidioa is Greek
for "club"). Familiar gilled mushrooms bear these spore-releasing
basidia on their lower surfaces. Basidiomycotes include everything
 Flesh of the Earth 177

fronr the corunon supermarket mushroom (Agaricus) to the Amanita

ulrosa (destroying angel) and its cousin, Amanita caesarea (a favorite
of the Roman emperor Claudius). Among their numbers are to be
found giant puffballs (up to nvo feet in diameter), earth stars (look-
ing Like tiny breasts decorated with the leaves of a jester's cap), smuts,
rusts, and jelly fungi (see plate r9b).
A fourth group, the deuteromycotes, consists of molds that form
neither basidia nor asci. The deuteromycotes probably lost this abil-
iry when they lost sex. But they are nevertheless reproductive wiz-
ards, capable of the ceaseless production of airborne propagules.
These organisnrs, also called conidial fungi or Fungi ImperGcti, re-
produce by conidia, thin-walled cells that break offfrom the tips of
ordinary hyphae. Others lack special reproductive structures; any
part of the body, any hyphal thread or mycelial mass may break off
to reproduce.
The fifth phylum of fungi consists of low-lyingphotosymthesizers-
the lichens. Lichens are one of the most striking examples of sym-
biosis. They are also among the most successful fungi. Like the bac-
terial merger that led to algae, the lichen is a combination of fungus
and alga (or sometimes, cyanobacterium). The result: an altogether
new life form that takes advantage of the alga's abiliry to make its
own food and the fungus's abiliry to store water and fend offthe
elements.

CROSS-KINGDOM ALLIANCES

All lichens-estimated at some 25,ooo different kinds-result from

cross-kingdom couplings between fungus and either green alga or
blue-green bacterium. Many lichens even harbor both rypes of pho-
tosynthesizers at once. Dwelling on bark and clinging to gravestones,
sheer cliffs, and other sunlit places unavailable to less enterprising
organisms, lichens have created a cozy niche for themselves. As they
grow, they slowly turn solid rock inside out, into crumbling soil and
living earth.
 178 | wnut ts Life?

Divided, the fungal gray and photosynthetic green components

of a lichen look nothing like one another. Nor does either mem-
ber resemble their extraordinary composite. The result of symbio-
sis, far from being predictable by simple addition, is a noncumula-
tive surprise.
With so many differenr lichens. each representing a permanenr
tryst between fungus and photosynthetic life form, the phrase
"long-term relationship" takes on new meaning. When the photo-
synthetic partner of a lichen is removed, sugar excretion by this pig-
mented partner stops and does not begin again even if extracts of
the sugar-inducing fungus are reintroduced. Somehow, algae and
fungi sense each other's whole-body presence as they form an en-
terprising, compiex partnership that depends on the history of the
relationship. Like animal cells, the algal and fungal cells in a lichen
communicate metabolically. Unlike most animals, however, the size
and shape of any variefy of lichen is not precisely fixed and the ex-
tent of their tissue complexiry is limited to one or a Gw tissue lay-
ers. Lichens, however, surpass animals in their longeviry; individual
lichens may be four thousand years old.
The ultimate interkingdom alliance, although not yer a lichen,
may be the one that exists today in Antarctica. Sevenry percent of
Earth's fresh water exists in Antarctica, but the water there is tied
up as ice, and the relative humidity at these forsaken outposts rarely
exceeds thirfy percent. The few regions of Antarctica that are ice-
free are thus deserts-and they are the driest places on Earth. Yet in
the barren expanse of these cold deserts, endolithic fungi grow with
green algae inside the rocks. Feeding offthe algae with which they
loosely reside (and which obtain sunlight through the translucenr
rock crystals), endolithic fungi derive their water from rare but ad-
equate melting frost.
Life can evolve suddenly, by jumps, when separate parties unite.
Interkingdom alliances between fungi and algae produced lichens;
a similar alliance may have been crucial to the developmenr of the
first forests. Root growths called mycorrhizae result from the dual
 Flesh of the Earth 179

growth of fungi and plants. Mycorrhizae provide the autotrophic

plant partner with mineral nutrients, whereas the heterotrophic fun-
gal partner is supplied with photosynthate food.
Mycorrhizae-rounded, stubby, often colorful roots-are sym-
biotic, a dynamrc structure produced by plant andfunzus. More than
five thousand distinct mycorrhizal fungi have been discovered. Most
of the associated plants seem to depend on these symbioses for their
supply of soil phosphorus and nitrogen. Mycorrhizae look neither
like plant root hairs nor like the fungus's mycelial nerwork. They
are synergistic, emergent structures, crucial to recycling. A single
large tree may have a hundred different mycorrhizae, produced by
distinct fungi, living in its roots.
Plants and fungi joined forces from the very start of life on land.
Some of the oldest plant fossils in the world retain evidence of sym-
biotic fungi. Lacking leaves, branches, and twigs, the first land plants
were little more than upright green stalks. Peter Atsatt, botanist at
the LJniversity of California at lrvine, and Kris Pirozynski, mycol-
ogy researcher formerly at the Museum of Nature in Ottawa, both
contend that successful colonization of the land by the ancestors of
modern land plants would have been impossible without root fungi.
Today fungi are still synergistically interrwined in the roots of more
than 95 percent of plant species. The algal ancestors of plants may
not have been able to come ashore without nutrient-procuring
fungi. The primeval arboreal carpet, the first forest floor, appears to
have been created notjust by plants but by plants and fungi acting
together.
Kingdom Plantae is (and always has been) almost entirely terres-
trial. The algal ancestors of this kingdom, of course, emerged from
aqueous environments, but most of the descendants kept to the land.
Early plants on land had to overcome almost insurmountable odds.
Land then was mercilessly irradiated and depleted of the nitrogen
and phosphorus salts absolutely required for plant growth. More-
over, the land was and is an unreliable provider of that vital resource,
water. Until the Silurian, when plants and fungi began to occupy
 'r8o I
What ls Life?

FIGURE 16. A gall on a

twig of Quercuq oak tree. Galls,
"disease" structures, may rep-
resent symbiotic organs in an
early phase of development.
Such bulbous growths occur
when plant tissue interacts with
fungi, insects, and perhaps bac-
teria. lt has been theorized that
galls were the evolutionary pre-
decessors to the first fruits.

the land, cyanobacteria and many other kinds of bacteria had the
desolate continents to themselves.
Kris Pirozynski has hypothesized that fruits-whose colors,
flavors, and aromas still cast an aesthetic spell over our primate
brains-evolved by way of interferences from the fungi and animal
kingdoms. His hypothesis attempts to explain the gap in the fossil
record between the spread of flowering plants and the appearance
of fleshy fruits a good forry million years later. Pirozynski envisions
that the first fruit appeared when fungal genes were transferred into
plant chromosomal DNA. This is similar to what occurs in crown
gall disease. Galls are symbiotic tissues formed by insect, bacterial,
or fungal growth on plants (fig. 16). They are bloated, somerimes
 Flesh of the Earth 141

slighdy monstrous-looking tumors found mainly on shrubs and

trees-and some look remarkably like fruits.
Crown gall disease is caused in many plants by penetration of
Agrobacterium. Agrobacterium, akind of bacterium that lives in soil,
bears plasmids (short pieces of DNA) that can enter into the cells
within roots and stems of susceptible plants, bringing bacterial genes
into the plant's nuclei. Biotechnology firms use Agrobacterium to in-
troduce desirable genes into crop plants. Pirozynski speculates that
fungal genes may have infiltrated plants in similar fashion. That fun-
gal inGctions were important to the abrupt, relatively late appear-
ance of fruits in many different flowering plants in the Cretaceous
fossil record remains an intriguing hypothesis. That galls are exam-
ples of plant-fungal synergy, however, is an established fact.

UNDERBELLY OF THE BIOSPHERE

Fungi resemble animals in that, unable to produce food, they de-

pend for nutrition on the bounry of others. From an ecological
viewpoint, however, the rwo kingdoms differ markedly. Fungi are
indispensable to the formation of soil, breaking down intractable
rock. They help lay down the carpet of spreading life. They are the
underbelly of the biosphere.
'Without fungi, plants and eventually all animals would be starved
for phosphorus (part of the autopoietic imperative as an essential
component of RNA, DNA, and ATP). Fungi mediate the inter-
stices of the food web. The Arab scholars who classified fungi mid-
way berween the plant and mineral kingdoms had a point. When
fungi take over a body, its material nature is quickly revealed. Bod-
ies become carbon-rich humus. Fungi decompose corpses and Ged
on living tissue, such as the skin of sweaty feet. For more than 4oo
million years their spores have been settling and sending mycelial
ner'uvorks through a global smorgasbord of foodstuffs. Recycling the
dead, they are the garbage collectors of the biosphere.
Fungi break down bread, fruit, bark, insect exoskeletons, hair,
 182 What ls Life?

horns, cameralens mounting compound, film, skin, buildingbeams,

cotton, feathers, and the keratin of fingernails and scalps. Like por-
table cleanup crews, fungi are transported around the world by air-
borne spores. Almost nothing is exempt from their gastronomic min-
istrations. Indeed, their zeal for recycling is so great that many begin
even before an organism has died. In maladies such as athlete's foot,
jock itch, and ringworm, fungi get ahead of themselves in the job
of redistributing elements of the biosphere.
Whether growing on human epidermal cells, breaking down the
cellulosic fibers of cloth with cellulase enzymes to create mildew
or making green-gray spores as a penicillium mold begins its colo-
nization of grapefruit, fungi digest substances left over by others.
'What seems like "decay"
to us is, to fungi, the healthy growth of
new offspring. Without fungi and bacteria breaking down complex
macromolecules, the corpses of plants and animals would pile up,
thereby taking phosphorus and nitrogen out of circulation.
Fungi, on land, perform most of the waste management func-
tion in the biosphere. Unlike ordinary people, who have survived
as polluting nomads for generations, dumping and moving on, the
biosphere cannot siniply put its refuse out on the planetary curb.
On Earth, garbage does not go out, but around. Humans are only
now approaching the level of sanitary efliciency mastered 4oo mil-
lion years ago by the fungi: fungi do not simply remove garbage,
they recycle it. Supplementing the bacteria, fungi recycle carbon,
nitrogen, phosphorus, and the like; on a plant- and animal-domi-
nated continental landscape they expanded planetary autopoiesis to
dry land, changing Earth's surface forever.

HITCHHIKING FUNGI, COUNTERFEIT FLOWERS,

AND APHRODISIACS

As rapid recyclers fungi often send the rest of us mixed messages.

Mediating at the interstices where garbage becomes food and corpses
become fertfizer, fungi, with their repertoire of hallucinogens, tox-
 Flesh of the Earth I f Sa

ins, and antibiotics provoke, deceive, and simulate the nervous sys-
tems of animals.
Ever since amphibians and their descendants ambled onto land,
we animals have had to contend with fungi. Indeed, for mrllions of
years fungi and animals have coevolved. Our primate ancestors
dwelled in forests and tasted many foods. Some were poisonous, oth-
ers mind altering. When spores or hyphae, resisting digestion, can
pass through animal intestines, it may be advantageous to them to
be eaten. Those animals who find fungi delicious often offer them
a nonstop free ride to the soil. For their part, animals that detoxified
or vomited poisonous fungi survived to unwittingly disperse them.
As language evolved, social prohibitions against ingesting possi-
bly dangerous fungi developed. So did sacred rites involving their
use. The attempts of societies to try to rid themselves of drugs
deemed threatening is reminiscent of the body's autonomic response
to rid itself of certain fungal "foods." But due in part to the abil-
iry of certain species to produce mind-altering trips, fungi will never
be completely eliminated from the body politic. Fungi are an en-
trenched part of sentient life in the biosphere.
In their ancient capaciry as sanitary engineers fungi have evolved
some rather startling relationships with members of other kingdoms.
Phallus and Mutinus are penis-shaped stinkhorns of the order Phal-
lales, whose stench, reminiscent of decaying meat, attracts flies. The
flies that alight on them carry sticky fungal spores away on their legs.
Pilobolus crystallinus is akind of mating (or "kissing") mold whose
favorite habitat is horse dung, rich in undigested cellulose, nitro-
gen, and other nutrients passed up by animals. Indeed, dung is so
valuable to fungi llke Pilobolus and other mating molds that they have
evolved a clever stratagem to get there first: they arrange to be eaten,
lying in wait amrd blades of edible grass. Mature Pilobolus does not
linger in the dung. Its sporeheads absorb water from the feces, in-
ternally building up intense pressures. Tensed for action, the spore-
heads, unbranched structures t\jvo to four centimeters in height, aim
toward the light. When internal pressures exceed 7oo kilograms/cm2
 184 What ls Life?

the sporeheads pop out of the manure-and land several meters

away, in grazeable grass. Here the long jumpers, an inspiration to
the internationally acclaimed Pilobolus dance troupe, can be earen
by helpful, munching horses.
Fungi have evolved other ruses. One yellow rust fungus, a basid-
iomycote, invented the insect-fooling stratagem of mrmicking a
flower. After inGction the rust fungus induces rock cress (a plant in
the mustard family that grows in Colorado's mountain meadows)
to metamorphose and mimic the common yellow buttercup. Rock
cress has unspectacular, droopy flowers but, once fungally infected,
it elongates and makes a nectar-rich rosette-a rigid uplifted "flow-
erhead" that attracts would-be pollinators. Instead of pollen, the vis-
iting insects pick up spores from these counterfeit flowers.
More than forty fungi species, such as Panellus, a delicate capped
stalk, glow in the dark.
'Why
they luminesce is not known, bur a
good guess is that in doing so they atffact animals such as nema-
todes, tiny translucent worms. The worms eat the fungus, excret-
ing and spreading the indigestible sticky fungal spores.
As ancient pioneers of land, fungi work with more recent set-
tlers to propagate themselves and their olTspring. The extraordinary
willingness of fungi to perpetuate themselves by any means, and
through any ends, possibly climaxes in the evolution of a particu-
lar fungus-and-ant agricultural system. Ever loyal to the fungi they
cultivate for food, ants of the genus Atta evolved wheelbarrow-like
depressions in their backs for carrying spores, which they then fer-
tilize and feed with chomped-offbits of leaves, bark, and other plant
material. These insects farm fungal spores as if they were seeds, care-
fully weeding out debris that might spoil rheir underground gar-
dens. As with the fungal flower mimics, this cross-kingdom associ-
ation brings to mind science fiction stories of beings enslaved to the
dull task of propagating other beings. But in this instance, the ants,
like harvesters enjoying the fruits of their agricultural labor, are fed
for their efforts.
Human farmers have domesticated apples, bananas, and straw-
 Flesh of the Earth I r eS

berries to the point that the sexually produced seeds are now in-
fertile. To grow at all, the plants must be grafted or "vegetatively
propagated" by human hands. Sirnilarly, the ancestral sexuality of
the basidiomycote fungi has been suppressed by long association with
Atta ant farmers. In tending to their fungal gardens, ants have sim-
plified fungal reproduction such that basidiospore-bearing mush-
rooms, normally the outcome of fungal sex, no longer develop. In-
stead of mushrooms, bulbous structures called staphyla crop up as a
result of all the plowing and planting. Like miniature fields of grain,
rows of edible staphyla are harvested by the ant farmers as the ma-
jor source of food for the colony.
The supermarket fungus , Agaricus brunescens, is now grown in large
limestone caverns in Pennsylvania. It has been thoroughly domes-
ticated. ("lt's like any other business," quips one executive familiar
with commercial mushroom growing. "You keep 'em in the dark,
feed'em manure, and when they pop up you cut their heads off'")
To gather Perigord black truffies, species narne Tilber melanosporum,
proverbial pigs (now more often dogs) are used to sniffthe redolent
hillsides of Provence in southern France and of Umbria in the Ital-
ian countryside. Although no farmers have yet been able to culture
this delicacy, savr1, mushroom lovers scatter tru{fle spores on the
roots of oak seedlings before planting, just to raise the probability
of finding mature truffles later.
Even the larsest black truffies are less than three inches in diame-
ter and weigh under rwo ounces. Tiuffies contain alpha-androsterol,
a steroid that is also foundin the breath of male pigs and thus may
be the reason why sows were traditionally used to hunt the fungi.
This steroid compound has been isolated from men's perspiration
and the urine of women. It may be an ingredient not only in truffles
but in the natural perfumes attracting the sexes. Rare as they are,
trufHes would be even more so were it not for their abiliry to en-
tice the sensibilities of certain mammals. For thousands of years
truffies have been devoured by pip, dogs, squirrels, people, and other
mammals, who (inadvertently or not) are all quite hrppy to spread
 186 What ls Life?

the spores throughout the woodlands in their droppings. Who

knows? The mammalian habit of burying excrement-like bury-
ing the dead, of possible benefit to these odd beings-may be yet
another fungal machination.

HALLUCINOGENIC MUSHROOMS AND DIONYSIAN DELIGHTS

Tiuffies are not the only offerings made by Krngdom Mychota for
the refined human palate. 'We savor gorgonzola and other blue
cheeses, halCdigested by molds. Nevertheless, we turn up our noses
when other fungi invade rnilk products, declaring them "spoiled."
Isoburyric acid lends its characteristic odor both to vomit and to
some of the finest French cheeses.
Fungi are a diverse and wild bunch. Mushrooms convey some-
thing musfy and strange and are often bypassed by foraging humans
because of the difficulry of distinguishing edible from poisonous va-
rieties. Indeed, it is at the interstices of edibility and toxiciry that
fungi produce their greatest manifestation: visions.
ln the Rig-Veda, aHindu sacred text, mention is made of drink-
ing soma and of finding one-footed beings that live upside-down in
the shade. These beings may have been mushrooms consumed for
religious practices. The Eleusinian mysteries of ancient Greece also
may have included the ingestion of fleshlike hallucinogenic mush-
rooms, perhaps Amanita, as part of a secret, drug-taking ritual.
In his written dialogues Plato records that Socrates, who left no
writings of his own, gave the name pharmakon (both "remedy" and
"poison" in Greek) to writing. The metaphor concerns writing's
pharmaceutical nature: it expands memory, yet breeds dependency
on writing implements. Even as writing expanded the storehouse
of human knowledge, it weakened traditional storytelling skills.
Socrates' argument against writing resembles the modern argument
that television has a druglike effect on children and thus interferes
with their learning (presumably more imporrant) reading and writ-
ing skills. Or again, the comparison of writing to a drug is similar
 FIesh of the Earth I lez

to the notion that electronic calculators should be banned because

they prevent students from learning math for themselves. But hal-
lucinogenic fungi are much older temptations than are writing, tel-
evision, or handheld calculators. They worked their strange magic
on our fruit-foraging primate and pre-primate marnrnalian ances-
tors before anyone was in a condition to discuss it.
Because fungi feed on animal feces (and corpses), their interest
often lies in making a rapid tour through the animal digestive tract;
they entice animals to eat them, but the spores and even the hyphae
may resist digestion. Social mamrnals and birds serve fungi, trans-
porting spores in their feces, sneezes, warm feathers, and fur.
Hallucinogenic fungi, such as Psilocybes mexicana, put a strange
rwist on the ancient relationship befrveen mammals and fungi. The
Catholic rite of the Eucharist, the flesh of God as mere wafer, may
have originated in pre-Christian initiation ceremonies involving the
ingestion of hallucinogenic fungi. Investment banker Gordon Was-
son (r898-1986) even suggested that the "apple" Eve plucks from
the Tree of Knowledge in the biblical Garden of Eden is actually a
rnistranslation of the word for an hallucinogenic fungus. An inter-
nationally renowned pioneer in the field of ethnomycology, the
scholarly-Wasson and his wife, Valentina, sought mushrooms in re-
mote villages in Mexico during the mid-r95os. Their experiences
watihing and participating in the sacred mushroom rituals of the
Mazatec Indians were recorded in a 1957 issue o{ Life rnagaztne,
and inadvertently, by informing the general public that such fungi
edsted, helped spark the psychedelic movement of the r96os and
beyond.
'Wasson
coined the term "mycophilic" to distinguish those cul-
tures (notably in Russia, elsewhere in eastern Europe, Catalonia in
northeastern Spain and southwestern France, and most of Korea and
China) in which people are familiar and appreciative of fungi, from
"mycophobic," or fungi-dreading cultures. Fungal ambivalence
roughly parallels the divide along the former Iron Curtain. Save for
Michigan and.Wisconsin morel pickers and chanterelle fans of Wash-
 '1 88 What ls Life?

ington and Oregon, the average American ciry dweller remains a

borderline nrycophobe no more likely to appreciate members of rhe
Kingdom Mychota than to think of herself as a symbiotic permu-
tation of bacteria.
Fungi provide an enchanting ber,ry of products, a veritable Di-
onysian garden of delights. They make the alcohol in champagne,
wine, and beer; they raise bread and give it its flu1Iy texture; they
ripen Brie, Camembert, Tioyes, Thenay, and Vend6me cheeses; they
flavor soy sauce and miso; they grow in the woods as delicious
trulfles, gourmet girolles, chanterelles, and morels. Oyster mush-
rooms, the saffron mrlk cap (shown on frescoes at Pomperi), the fairy
ring mushroom (often dried and used as spice), the dangerous-look-
ing bluish purple blewitt (sold in markets in northern England), shi-
itake mushroorns (I-cntinus edodes), the enokitake or velvet stem
mushroom, the nameko, the high-priced pine truffie or matsutake,
and Boletus edulis Qtorcino to the Italians, Steinpilz to the Germans,
setd to the Spanish, cipe to the French) all may be eaten with Ra-
belaisian relish.

TRANSMIGRATORS OF MATTER

The Chinese elephant ear fungus, corrrnron in hot-and-sour soup,

is thought to prevent cancer and retard heart disease. Fungi indis-
putably produce antibiotics. Scotsman Alexander Fleming (r881-
I95 5) found quite by accident that a greenish mold, Penicillium, pre-
vents bacterial growth. Penicillin, the active ingredient produced by
this ascofungus, inhibits bacteria from making their cell walls, fight-
ing strep throat, bacterial pneumonia, and other once-untreatable
infections. Another mold, Tblypocladium, produces cyclosporine, one
of the least toxic and most effective inrmunosuppressants known,
used to prevent infections after organ transplants.
For every fungal evil-every refrigerator crisper invaded by a black
stain, every retching resulting from an incorrectly identified mush-
room, every peach gone the way of green fozz-there is an equal
or overcompensating fungal good. This book would not be possi-
 Flesh of the Earth 189

ble without fungi: nutrient scavengers fungi connect and feed the
as
roots of trees, providing the wood and pulp precursor to this page.
"Yes, happily language is a thing," writes the French literary theo-
rist Maurice Blanchot, "it is a written thing, a bit of bark, a sliver
of rock, a fragnent of clay in which the realiry of the earth con-
tinues to exist."+ Composed of dead and living things, a place where
dirt comes clean and where wastes are reborn as hyphae and spores,
the soil is largely a fungal phenomenon. Fungi are part of "the re-
aliry of the earth." They are the most resilient of land's eukaryotes,
able to thrive on great variery of compounds, break down com-
^
plex organic molecules, and inhabit seemingly inhospitable domains,
such as the inside of glaciers or the extraordinarily acid waters of
Rio Tinto in southwestern Spain.
The lifesryle of fungi alerts us to the arbitrariness of our notions
of individuality. Growing without discrete borders, fungi mate
pronriscuously with those of many complementary genders. Their
tiny mated threads produce distinguishing organs of sexualiry rec-
ognized as mushrooms, puflballs, shelf fungi, and the like. Many
team up with plants, algae, or cyanobacteria to form composite or-
ganic beings: the nrycorrhizae or the lichens. Planetary physiology-
Gaia-is the result of interaction of innumerable beings, including
the fungal networks. Gaia is symbiosis seen from space. Fungi cy-
cle matter and turn waste into usable nutrients within a symbioti-
cally integrated biosphere.
Any organism that appears or species that evolves at first has a
chance. But to persist, life forms must survive not on their own but
within a global environment. They become integrated, or they die
away. As the poetTlkami attests, people tend to consider fungi freaks
of nature, frightening and expendable. And yet, from a planetary
perspective, fungi proved themselves long ago. For 4oo million years,
'With spores
from tropics to poles, they have been untiring recyclers.
riding the winds or hitchhiking on kin from other kingdoms, fungi
have spread themselves about the globe, adventurous pioneers and
desiccation resistors par excellence.
When animals die, fungi give them a natural grave. By way of
 190 What ls Life?

fungi, corpses come to fortifii grasses and trees. Sawed and crushed,
their cellulose fibers become papeg books, immortal words, or words
to be recycled into more words. Reminiscent of the Eastern doc-
trine of the transmigration of souls, fungi are the transnigrators of
matter.
Life creates. The global autopoietic system, Gaia, spins off crea-
tures increasingly strange. For a while at least, even for millions of
years, the global environment will tolerate bizarre sports, rapidly
spreading pioneers, opportunistic monsters. But in the long run or-
ganic beings confront the limits of their own multiplication. They
survive not alone but within a context of global life. Chomping lo-
cust swarms devour monocrops; guano birds defecate salts, moving
phosphate and nitrate from sea to land. Rapid reproducers begin as
inGstations or infections but are tamed. Any rampant planetary pop-
ulation, any out-of-control "tumor," finds its economy. All grow-
ing populations integrate into the working biosphere or they be-
come extinct.
In English the word "fungus" is virtually synonymous with an
unwanted, surgically expendable outgrowth. Such a meaning nright
apply better to the hoarding, rnaterialistic species we have become
than to the organisms that nobly serve as biospheric undertakers,
investing animal waste with life and turning corpses to soil.
Fungal spores may once have shared the air over land mostly with
the propagules of far more ancient cyanobacteria and bacilli. In the
long run, however, they were to intimately share the atmosphere
with ameba and algal cysts, bacterial spores, fern spores, flowering
plant pollen and seeds, and with flying insects, birds, and bats. They
distributed themselves by ascospores, conidiospores, basidiospores,
activated dry yeast, and as lichen propagules: soredia and iscedia. Not
only colonizers, they became powerfully selected degraders, recy-
clers, and agents of planetary redistribution.
Those of us who succumb to prograruned death and whose re-
mains are neither scavenged nor flamed go to a fungal underworld.
The chemicals of our bodies are returned to the earth. Fungi keep
 Flesh of the Earth I lS't

life's compounds recycling. The global workings of Kingdom My-

chota run as smoothly as a kidney or liver. We, with our nomadic
past, are still adjusting to the idea that in a closed system the fruits
of our labor and loins cannot accumulate endlessly. They must be
distributed, returned to the system whence they came. This is a
difiicult lesson: the matter of our bodies, our possessions, our wealth
is not ours. It belongs to Earth, to the biosphere and, whether we
like it or not, that is where it is headed, again and again. Fungi help
get it there.

SO, WHAT lS LIFE? Life is a net'vvork of cross-kingdom alliances,

of which Kingdom Mychota is a willing and crafty participant. Life
is an orgy of attractions, from the trickery of counterfeit "flowers"
to the strange allure of truffles and difiicult-to-swallow hallucino-
gens. As fungi, life seeks out the underworld of soil and rot no less
than the sunny vistas overwhelmed by photosynthetic beings. Life
is self-renewing and fungi, as recyclers, help keep the entire plan-
etary surface brimrning with life. Transmigrating matter, molds and
mycelia have found their calling. Creating and destroying, attract-
ing and repelling, undertaking and overturning, they are part and
parcel of terrafirma.
 THE TRANSMUTATION
OF SUNLIGHT

Tygerl Tyger! burning bright

ln the forests of the night,
What immortal hand or eye
Could frame thy fearful symmetry?
WILLIAM BLAKE

lf one has the patience, and the courage, to read my book, one
will seethat it contains studies conducted according to the rules
of a reason that does not relent . . . but one will also find in it
this affirmation that the sexual act is in time what the tiger is
in space. The comparison follows from considerations of energy
economy that leave no room for poetic fantasy, but it requires
thinking on a level with a play of forces that runs counter to
ordinary calculations, a play of forces based on the laws that
govern us. ln shod, the perspectives where such truths appear
are those in which more general propositions reveal their mean-
ing, propositions according to which it is not necessity but its
contrary, " luxury," that presents living matter and humankind
with their fundamental problems. . . freedom of mind . . .
issues from the global resources of life, a freedom for which,
instantly everything is resolved, everything is rich.
CEORCES BATAILLE

The action of solar radiation on earth-processes provides a

precise basis for viewing the biosphere as both a terrestrial
and a cosmic mechanism. The sun has completely transformed
the face of the Earth by penetrating the biosphere, which has
changed the history and destiny of our planet by converting
rays from the sun into new and varied forms of energy. At the
same time, the biosphere is largely the product of this radiation.

VLADIMIR VERNADSKY

193
 't94 What ls Life?

GREEN FIRE

The ultimate source for all life's energy, growth, and behavior is the
sun. Burning like a cool green fire, photosynthetic beings rransmute
sunlight into themselves (see plate zo). Protoctists (coccolithophorids,
diatoms, seaweeds) are the main transmutors in the sea; plants the
main ones on land.
Plants represent a high point in bacterial coevolution. They have
raised the biosphere to a higher dimension-up to roo meters from
the soil surface. Yet they are newcomers to the photosynthetic guild.
Plants have only dwelt onEarth for the last 45o million years. Evolv-
ing from algae, plants-almost exclusively land beings-went on to
green the continents.
The blue whale, 26 meters long and weighing r 8o,ooo kilograms,
is the most massive animal ever to have lived, heavier by far than the
largest dinosaurs. Nonetheless, next to behemoths of the plant
world-such as the giant sequoia, which can weigh rwo rnillion
kilograms-even whales are 1ight. One clone of the quaking aspen,
Populus tremuloides, is estimated to contain forfy-seven thousand
trunks. Nominated by Universiry of Colorado biologist Jeffry
Milton as perhaps the biggest individual organism on the planet, this
dispersed but connected tree covers forfy-three hectares in Utah. It
is estimated to weigh six million kilograms (plate zr).
Books come from plants. So do boardwalks, oak desks, hashish,
cotton shirts, chewing gum, coal, myrrh, clapboard houses, choco-
late. Plants are the source of morphine, codeine, heroin, and other
drugs similar to endorphins-pleasure-giving chemicals produced
naturally in the mamrnalian body. Bark of Salix, the willow clan,
gives us salicylic acid, aspirin; other plants make not only analgesics
but astringents, antifungals, antispasmodics, pigments, caustics, car-
diovascular agents, expectorants, diuretics, fumigants, hemostatics,
insect repellents and toxins, perfumes, and anti-asthmatics.
Plants have been such a deeply embedded part of the human en-
vironment that we now hardly notice them. (Jnless a bouquet of
 The Transmutation of Sunlight 195

long-stem roses or box of chocolates arrives on the doorstep, plants

are taken for granted. Even then, meaning or mood is evoked by
the plant product as a symbol rather than the plant itself.
Plant life presents us with an extraordinary richness of sights,
smells, and tastes. The seasonal bursts of fragrant flowers have a
beneficial psychological effect for dwellers outside the tropics; the
mere sight of hills of undulating grass can produce serenity.
Of the nine recognized phyla of plants, only one has flowers. But
that one phylum is so diverse that it is thought to account for more
than half of all plant species. A full documentation of all species in
the three hundred famrlies of flowering plants would be such a mon-
umental task that it has never been undertaken. "Such a listing,"
writes botanist Frits Went (r9o3-r99o), "would have to describe
about a quarter million known plants; to compile it, all the taxo-
nomic botanists in the world would have to work together for years
and years, and the finished product would have perhaps half a mil-
lion pages, enough to cover a whole wall in a library."l
Plants were not, however, the first form of "green fire" on land.
Fifry miles southwest of Las Vegas, an 8oo-million-year-old fossil
soil is preserved in rock. The carbon content indicates some kind
of ancient photosynthetic life. From another spot in the American
Southwest, eighty miles northeast of Phoenix, Arizona, even older
samples of fossil soils were collected, corroborating a hypothesis by
Susan Campbell and Stj epko Golubic of Boston LJniversity that pho-
tosynthesis on land began in cyanobacterial form I,2oo million years
ago or earlier. Paleontologist Robert Horodyski of Tulane fJniver-
sity in New Orleans and geologist L. Paul Knauth at Arizona State
lJniversiry in Tempe contend that land was abundantly covered by
photosynthetic microbes in the late Proterozoic eon.2
Not until the Silurian appearance of true plants, with their alter-
nation of spore propagules (formed by meiosis) and gametes in sex
organs (that fuse to make embryos), was there a full-fledged escape'
probably at first seasonal, from the algal necessiry of dwelling in wa-
ter (see plate zz) . Freeing itself of water, life on land evolved inter-
 196 What ts Life?

nal means of support and grew up on land. Land plants made a

water-pressure system of structural support from the cellulose mol-
ecules found in bacteria and algae. Later, they evolved a stronger
substance that, when combined with cellulose, would remain elas-
tic but strong and supportive, even in dry conditions. This subsrance,
lignin, is the chemically complex polyphenol that gives woody
plants their woodiness. With lignin, the biosphere began its verti-
cal climb, extending life's realm to another, third dimension over
the land. Biologist Jennifer Robinson has suggested that the great
piles of coal left in the Earth's crust owe their existence to the lag
after plants had invented lignin but before fungi had evolved means
of decomposing it.
Just like other life forms, planrs came from microbial predeces-
sors.Their heritage is of photosynthesis, but they need not be pho-
tosynthetic. Sorne plants, even those with leaves and fruit, have aban-
doned the way of green fire, no longer photosynthesizing. Like those
eyeless, subterranean mole rats that no longer see because they do
not need to, some white plants have outgrorvn their dependence on
direct sunlight. For example, Epi-fapus (beech drops) and Monotropa
(Indian pipes) absorb their mineral nutrienrs through the subvisible
fungal threads, intertwined in their benefacrors' roor hairs, that ex-
tend to those of neighboring green forest trees. From the white plant
to the tree, roots (like animal organs) are connected by fine, long
nerve-like threads of the fungus's body.
The distinguishing feature of plants is not photosynthesis per se
but that they all grow from spores at one stage in their life cycle and
from embryos in another. The plant embryo, found deep in the ma-
ternal tissue, is the diploid product of sexuai union. While it is, like
an animal embryo, formed from the fusion of a male propagule with
an egg held in a female organ (rhe "archegonium" in plants), nei-
ther the egg nor the sperm of a plant is produced by meiosis. plant
embryos form when male pollen tubes or swimrning spernl pene-
trate tiny female plants or the female parts of hermaphroditic plants.
These tiny female plants are haploid and grow from haploid spores
 The Transmutation of Sunlight 197

buried in the mother's haploid tissue. The mature plant that grows
from the ernbryo does not make gametes (as mature animals do);
rather, it makes, through meiosis in its diploid body, spores. The
spores grow into either male or female haploid plantlets that make
ganletes without meiosis (fig. tZ).
Plants are sexual beings. Sexual coupling is the act, and the em-
bryo is the structure, that distinguishes them from algae and mem-
bers of other phyla (lichens, for example) that have sometimes been
misrepresented as "plants." Plant sex, however, differs from that of
animals: although Grtilization of plant egg and sperm nuclei makes
embryos, plant meiosis does not make eggs and sperm. Meiosis
makes spores. Spores grow into plantlets, each with only a single
set of chromosomes. These plantlets, called gametophytes, can
grow-unlike an unfused animal egg or sperm. The gametophyte
grows by mitotic division of its cells, which carry only a single set
of chromosomes.
In cone-bearing and flowering plants the gametophyte (either
male or female) is just a tiny structure that is not free-living. The
gametophyte fornm and spends its life entirely within a cone or
flower of the plant that meiotically produced it. Plantlets make their
sex organs and their gametes by mitotic cell division. Because they
begin with only one set of chromosomes, these cells do not have
to change their chromosome numbers to produce sperm or eggs.
When the rnating cells fuse, doubleness is reestablished and the cy-
cle begins again.
Evolutionarily speaking, however, the failure of the gametophyte
to "leave home," so to speak, is a recent condition. Plants of an older
lineage, ferns for example, cycle through an alternation of genera-
tions in which the small bodies of those with a single set of chro-
mosomes and the large ones with rwo sets are physically unconnected
and strikingly different in form.
Overall, as sexual embryo-forms, plants and animals are more
alike than either are like the three other kingdoms ftacteria, protoc-
tists, and fungi). Animals are, however, diploids with a single-cell
 male flower petals

.]_ embryo sac

ovary: cross
,u/ /
section of
/1 female flower
anther

female sPerm nuclei

flower

egg cell
nucleus

male pollen
tube with
sperm cells

fertilization of
female by
flower pollen

acorn
(seed in fruit)

FIcURE 17. Differ"ent stages in the sexual life hlstory of an oak tree, Quercus. Counterclockwise
from bottom: oak leaves with two mature fruits that contain the seeds (acorns). The flower shown
at center left is magnified and its walls (the ovary) are cut away to reveal the eight nuclei of the
double fertilization of angiosperms (center). The pollen tube has penetrated the embryo sac and
let loose three small male nuclei. One will fertilize the egg nucleus (bottom center) and one will
fertilize two of the larger female nuclei to form triploid (3-chromosome set) tissue which will nourish
the embryo-hence "double fertilization. " The doublet anthers that produce the pollen are shown
on their stalks at the upper left. At the upper right a germinated pollen grain that has formed a
pollen tube is seen on its way into the ovary of a bisected oak flower.
 The Transmutation of Sunlight 't99

haploid phase, whereas plants have a multicellular haploid phase-

and, unlike the cells of animals, all plant cells contain rernnant cyano-
bacteria, plastids.

THE ACCURSED SHARE

In the enigmatic epigraph to this chapter, Georges Bataille (r8Sl-

196z) links the tiger with that initiating point of the mammalian liG
history, sexual intercourse. He assures us, moreover, that his compar-
ison is rational. It is. The entire unfolding of evolution is a response
to an unexportable excess, a growing surplus of sun-derived energy.
Both the sex act and the tiger are complexities of the biosphere.
While coitus is a behavior and the tiger a being, together they
represent rr,vo fates of plants' prodigious reserves. The tiger is poised
atop a pyramrd of global nutrition whose base is the sun. Even at
rest, the tiger represents life's nutritional edge, its carnivorous lirnit.
The tiger, "burning bright" in Blake's memorable phrase, represents
the funneling of solar radiation into a highly specific and potentially
terrifying form. Coitus employs sun- and plant-derived wealth as
animals expend energy to make more of themselves. Bataille fur-
ther argues that classical economics is rnistaken: the general econ-
omy is not human but solar. Sun-produced food, fiber, coal, and
oil-carbon- and energy-rich reserves-are the living foundation
not only for bustling animal life but for industry, technology, and
the wealth of nations.
The economy comes from photosynthetic life and the sun. Pho-
tosynthesizers use solar radiation to produce the cold hard cash of
the biosphere. Heat is dissipated, degraded energy lost to space as
primordial wealth accumulates. Colorful photosynthetic bacteria,
protoctists, and plants the world over produce and "save." Eating
them, consumers may "spend" gathered photosynthetic energy
through metabolic activities or anabolically (and temporarily) store
it in their herbivorous or predatory tissues. Primordial wealth may
also end up in long-term storage (or be lost outrighQ when con-
sumers die and are buried without decay.
 200 What ls Life?

Spending has always been a critical problem for life. Greed comes
easily within a biosphere whose constituency triumphs as a func-
tion of the ability to amass the wealth of photosynthesis. Bataille's
tiger mercilessly hunts the leaf-eating deer. North Americans now
fell plants to print paper money with colored fibers-or subrnit such
bills in return for the striped pelt of that endangered mammal. Pho-
tosynthesis creates excess, surplus, a reserve of matter and energy
whose uses are as numberless as life is creative.
Bataille perceived that the character of a particular sociery is de-
terrnined less by its needs than by its excesses. Wealth creates free-
dom in both biological and cultural realms. A nostalgia for old Eu-
rope, a respect for native American restraint, an admiration for the
opulence of Egypt-these are sentiments based implicitly on the
recognition that a culture is determined by how its members choose
to spend or accumulate its excess. Rome makes its coliseum and basil-
icas, America its McDonald's and Disneyland, Egypt its sphinx-
guarded pyramids.
In the United States politicians grapple with tax collection, deficit
and debt reduction, and public spending. The government prints
money that banks lend without having or touching. Stocks, bonds,
certificates of deposit, of
precious metals, and other instruments
finance are owned by investors. But what does it mean "to own"?
Humaniry does not own what it spends; ownership rests with the
biosphere. Checks, credit cards, paper nloney, and stock certificates
are all symbols of a wealth whose source lies beyond technological
humanity's means of production. The monetary economy attempts
to arrest the solar flux of Earth's economy. Money symbolizes the
conversion of photosynthesis, life's energy, into something else-
something that can be controlled, manipulated, and hoarded by hu-
mans. Perhaps it is no coincidence that in the United States money
is green.
The fact remains that without plants the vast majoriry of animals
would starve. Indeed, even with luxuriant plant growth humans and
all other animals are destined to die. The grave is a great leveler, and
 The Transmutation of Sunlight 201

a good reminder that we are owned by what we own. All of us from

street sweeper to billionaire pay our dues. The elements of our bod-
ies return to the biosphere whence they came. In the restricted econ-
omy of human arrogance and fantasy, individuals may amass great
wealth and power. But in the solar economy of biological reality
each and every one of us is traded away to make room for the next
generation. On loan, the carbon, hydrogen, and nitrogen of our
bodies must be returned to the biospheric bank.
A biosphere differs from an organism in that it is essentially closed
to influx and egress of materials. Although supplies of carbon us-
able by life arrived with meteors and comets that penetrated Earth's
atneosphere, especially before life took hold, this external source of
material today is insignificant. Unlike an organism, eating and ex-
creting, the biosphere has become self-contained. Its materials are
limited. Chemically transformed, they are used over and over, but
they are never used up. The luxurious surplus of edible and usable
compounds produced by photosynthesis leads to scavengers and
predators, organisms killing and eating or cleaning up to survive and
grow. The limited material reserves of the biosphere constrain the
amount of solar rays that can be transformed into green life.
Overall, photosynthetic activiry creates a surplus of energy-rich
matter that can be hoarded, eaten for growth, or outright squan-
dered. The great planetary riches are there for the taking, replen-
ished by the lively conversion of solar energy. It is an understand-
able but impossible wish to preserve the planet in its "original" state.
The pristine nature to which some wish to return is not eternal but
rather the green world that supported our ancestors so beautifully
that they overpopulated it. Moreover, human spoilage of the lush
environments that nurtured us is not evidence of any singular abil-
iry to imperil all life on Earth. No single species in the past has ever
threatened all the others. Any tendency of one kind to overgrow
and despoil was kept in check by all the rest. The essence of "nat-
ural selection" is that unstoppable tendencies of one population to
grow to the point of environmental degradation will be halted by
 202 What ls Life?

the growth of others. Human population expansion plays by the

same rules: the degraded environment breeds morbidiry high mor-
taliry and ultimately even extinction.
Our evolution has unearthed hoarded organic treasures, such as
coal and oil to power cars and heat homes. Wealth in the biosphere
ultimately comes from the sun. Organisms die, populations decline,
and species become extinct. But the biosphere gets richer. Human
burning of fossil fuels, for example, is exploited by plant life. Plants
incorporate carbon dioxide released from this burning into their
bodies. This is not to say that the current industrial mode of human
living may not be dangerous or lead to increases in global temper-
ature. Rather, the conversion to waste of a surplus by one life form
has biospheric precedents: far from impoverishing the planet, the
waste of one may, in fact, create more wealth for another.
In the strange solar economy individuals die, returning their bod-
ily wares to biospheric circulation. Chemicals used in bodies are not
lost. AII organisms confront the combined difiiculry and temptation
of making use of that persistent photosynthetically derived excess
to which Bataille gave the name "the accursed share."

ANCIENT ROOTS

The first plants were probably like today's bryophytes. Mosses, liv-
erworts, hornworts, and their kin lack the vertical stature of other
phyla, owing to the absence of any system for fluid transport and
hydrostatic support. Little more than masses of green cells favoring
moist surfaces, bryophytes then and now lack leaves, roots, and seeds.

At the end of the Ordovician period the land surfaces were coarse,
populated by low-lying cyanobacteria and soil algae but no plants.
Where water was dependable, in rivers and lakes and along the bor-
ders of the sea, cyanobacterial mats became thick. In drier locales
a sinuous, tough binding of blackish green soil particles and angu-
lar bits of rock covered the land. A modern analog of such terres-
trial life prior to plants are the desert crusts of lJtah, the Gobi Desert,
 The Transmutation of Sunlight I ZOa

and the fields of lraq. These crusts consist of cyano- and other bac-
teria, occasional algae, and fungi-all of which are quick to begin
the green fire of photosynthesis (or revert to a quiescent state) when
given moisture.
Some modern green algae ("ctrlorophytes"), especially the chaeto-
phorales, have been proposed as sirnilar to the ancestors of plants.
Their chloroplasts contain chlorophylls a and b-the same pigments
found in the chloroplasts of plants. Like the sperm tails of mosses
and ferns, rwo quite different sorts of plants, motile chaetophora-
lean algal cells bear rwo undulipodia. These green cells have inter-
cellular connections, plasmodesmata, that resemble the perforations
through the cell walls of plants. Animal cells join by strenghening
contacts utter\ unlike the perforations of plasmodesmata of algae
and plants. The details of their mitotic cell divisions and their walls
made of rypical plant cellulose suggest that certain chlorophytes, such
as the modern filamentous green alga Klebsorbmidium, resemble the
imagined ancestors to plants.
Today ferns, ranging from less than three centimeters to over
nvenry meters in height, still reproduce by aquatic methods of egg
and swimming sperm, which they shed into nearby puddles. Even
so-called higher plants, such as the Cinkgo (a showy tree with fan-
shaped leaves and stinking cherry-like cones, indigenous to steep
slopes in eastern China), bear their ancient heritage in the form of
undulipodiated sperm. The many tails of a single sperm in cross-
section are undulipodia with the same 9Q)+z symmetry of rnicro-
tubule arrangement that is found in motiliry structures from algal
swim tails and Paramecium ctlia to bull sperm and the fine hair cells
of the human lung.
The oldest well-preserved plant fossils are from black cherts at a
quarry in Rhynie, a hamlet in Scotland. Geologists believe that the
Rhynie fossils owe their superb preservation to periodic flooding
from a nearby, silica-rich spring. The fossil plants, such as Rhynia,
bear swollen roots, suggesting that fungi were already symbiotic with
plant roots 4oo million years ago.
 204 What ls Life?

A form suggesting the most ancient members of the latest king-

dom is still living today. This is Psilotum nudum, an obscure plant
dwelling in greenhouses and in Florida, the Pacific Islands, and other
sunny climes. Psilotum is a system of stems, a mere bundle of green
growing sticks. It releases spores into the air that produce sperm,
which can swim in most soil films or across puddles. After fertil-
izationthe embryo gives rise to new shoot growth. Like a bryophyte,
Psilotum lacks roots and seeds. But unlike a bryophyte, it has a vas-
cular system and stands upright. Lacking leaves, photosynthesizing
along its stems, it may resemble the earliest forms of plant life.
Modern nlosses and liverworts also suggest the shapes of prirni-
tive greenery. These bryophytes overcame the algal dependency on
a fluid surround by bringing water to land with them. The mere ex-
istence of leaves in mosses and liver-worts gave them a big edge over
the ancient psilophytes which, ltke Psilotum, were lirnited in their
abiliry to gather light. But, unlike vascular plants, the bryophytes never
evolved structural support. Bryophytes to this day never grow more
than few centimeters high. They are vulnerable to the tricks of vas-
a

cular plants that can easily overtop them, robbing them of sunlight.
Although no one is sure and the fossil evidence is scanry many
botanists believe that the simpler, more aquatic bryophytes evolved
earlier than did the structurally more complex and dryness-resistant
plants. Bryophytes are soft-bodied; their fossil record is decidedly
poor. Modern bryophytes are utterly dependent on surface waters;
they have no roots to scavenge for water down into the soil. But
they are by no means fragile creatures limited to swamps, pond
edges, river rocks, and waterfalls. Some live in areas of seasonal mois-
ture, growing mainly during the wet season. Others, notably the
ingenious sphagnum moss, are the sponges of the land. They are
world-class water scavengers, capable of holding up to a thousand
times their own body weight in water, storing it for dry times. A
mound of sphagnum moss, moreover, employs the dead in the task
of water retention. Only the surface of the mound is alive, but the
moss corpses in the interior and lower reaches retain water for their
descendants.
 The Transmutation of Sunlight 2O5

Most of plant diversification occurred in the vascular plants with

tough bodies and conductive tissue. These organisms grew up, lit-
erally. Horsetails, for example, were among the first organisms to
tower into the air. Modern horsetails, called "scouring rushes," have
silica in their photosynthetic stems. They were used by European
settlers of North America to scrub pots and pans. But these silica-
stiffened organisms grew much larger in the past than do their de-
scendants today. Ancient horsetails in primeval forests, in the De-
vonian period 4ro rnillion years ago, stretched up to fourteen meters
high.

PRIMEVAL TREES

Rhynia-type plants seem to have evolved into many extant and now-
extinct forms. The ancestral vascular form probably gave rise to pro-
gymnosperms-an extinct lineage that branched off in one direc-
tion to become tropical seed Grns, which themselves later gave rise
to the flowering plants. Another branch became the conifers that
brachiosaurs dined upon and that lived on to survive meteor im-
pacts and ice ages. The early Rhynia-rype vascular plants also diverged
to become ginkgos, spore-releasing ferns, horsetails, and Psilotum.
The branching talents of the original stem-maker thus came to en-
shrub and enforest the world.
A huge group of vascular plants, as important as the dinosaurs to
the aninral kingdom, have gone extinct. Known as the cycadofili-
cales or seed ferns, none of these trees-which looked like over-
grown pineapples-are alive today. They were not ferns at all. Un-
like modern ferns, they made conspicuously large seeds. These seeds
(not directly related to modern seeds) were a major evolutionary
innovation. Seeds can wait through a drought or cold spell. They
can survive a lack of light. Seeds were as crucial to the dispersion
of plants as water-tight eggs were for the great diversification of
reptiles.
Possibly the first plants to produce seeds, the cycadofilicales
abounded 34J to zz5 mrllion years ago, before any dinosaurs. They
 206 What ls Life?

were the makers of the earliest forests. Leaves of the genus G/os-
sopteris (Greek for "tongue-1eaf") are corunon fossils in rocks de-
posited in the southern supercontinent of Gondwanaland (see plate
z3). Exposed to powerful tectonic forces, Gondwanaland cracked
and the pieces drifted apart on continental plates zoo million years
ago. Those pieces are now called South America, Africa, Australia,
India, and Antarctica. Glossopteris, like more than 99 percent of the
plant and animal species recorded in the fossil record, is extinct.
Neither Clossopteris nor any of its once-successful relatives sur-
vived to inhabit the southern continents Gondwanaland has become.
But, once upon a time, greening the world for over roo rnillion
years, forests of seed ferns swayed in warm winds from the south-
ern reaches of Gondwanaland to the tropics of Laurasia, the north-
ern supercontinent. Now, after rz5 million years, the forests of
Gondwanaland exist as a semipetrified, energy-rich plant refuse:
coal.
By the end of the Devonian period and the start of the aptly
named Carboniferous (Mississippian and Pennsylvanian) 36o mrllion
years ago, Earth was forested. Whether from Rhode Island, Edin-
burgh, or western Pennsylvania, coal of this age is replete with re-
mains of leathery leaves, thick roots, and scaly bark. In the base-
ment of the Biological Laboratories at Harvard Universiry are "coal
balls" that were hauled away from their sites of origin in Illinois and
Kansas. Many are taller and, because they are spherical, stouter than
a man. Chopping through them, or peeling olf their surfaces with
acid-treated acetate, reveals the source ofancient plant tissue: leaves,
bark, roots, and flowerless sex organs hardly the worse for z9o mil-
lion years of burial.
Measured by genera and higher taxa lost, the mass extinctions of
the Permo-Tiiassic 245 million years ago were far more devastating
than the better-known end-of-Cretaceous event that extinguished
all the dinosaurs. A major factor in the Permo-Tiiassic extinctions
may have been expansion of glaciers or a long period of profound
cold-perhaps itself generated by a comet or meteor impact that
 The Transmutation of Sunlight 207

darkened the skies with debris sent into orbit. Seed ferns were trop-
ical plants. The seedlings of seed ferns and the trees themselves were
vulnerable to bitter cold. Before all the seed ferns became extinct,
however, at least one of their ancestors gave rise to plants that could
withstand freezing temperatures-the conifers.
Conifer fossils are older than those of flowering plants. Fossil seeds
of conifers are detectable as raised portions on the underside of female
cone scales. Spruce, cedaq pine, and many other cone-bearing trees
and shrubs alive today remain green all year long. So did many of
their ancestors, adept at surviving arid, wintry conditions. The pol-
len of conifers is wind-borne. Fertilization of conifers leads to for-
mation of seeds in the shelter of female cones (see plate z4). This
change, from the ancestral method of releasing delicate water-borne
sperm or short-lived spores such as those shed from the underside
of Grn fronds and destined to make tiny gametophyte plantlets, per-
nritted evergreen conifers to dorninate lands of seasonal ice and snow
or aridiry as they do to this day.

FLORAL PERSUASION

In contrast to all cone-bearing, naked-seed plants, the flowering

plants have encased seeds-the result of the growth of the flower's
ovaries into fruit. More than a quarter mrllion species of flowering
plants inhabit Earth. The seeds of these angiosperms bear embryos
and the fernale organs transform to encase them.
Humans have a special relationship with angiosperms. Our pri-
mate ancestors lived among African flowering trees and fed, in part,
on fruits-which had evolved luscious colors, arresting aromas, and
other tempting qualities insofar as such qualities seduced us into be-
coming involved in their reproduction. Mamrnals dispersed the en-
cased seeds and, by defecating, enriched the soil where angiosperms
would sprout. Ancestors more closely resembling modern humans
no longer lived in trees, but they kept their nimble hands and binoc-
ular vision while living in a new landscape that was another cross-
 208 What ls Life?

kingdom creation. The grassland savannas were the workings of an-

giosperms that invented a way to grow from the base rather than
their tips. Savannas were equally the creation of the large herbivores,
whose grazing killed tip-growing forbs and young rrees, thus "nat-
urally selecting" the grasses. Finally, the savanna would have been
impossible without its recyclers: protoctists and bacteria which di-
gested cellulose in enlarged fore- and hindguts coevolved in the large
mamnralian grazers.
Even today our species has a special relationship with angiosperms.
Angiosperm grains, fruits, stems, leaves, and roots are our primary
foods-directly, or indirectly by way of domesticated livestock. (The
only exceptions are intensive fishing comrnunities.) We surround
ourselves with furniture that is often made from the lignin of for-
est angiosperms. Angiosperms have taught us pleasure in nurturing
them in flower gardens. Even the image of a branching tree to ex-
plain evolutionary phylogeny is easy to understand in part because
of our ancient familiariry with the growth patterns of flowering
trees.
Charles Darwin called the origin of the flower an "abominable
mystery" (see plate z5). Beautifully fossilized flowers and seeds in-
dicate that flowering plants appeared in the middle latitudes of the
Northern Hemisphere at least by n4 million years ago in the mid-
Cretaceous. Thus, within sixry million years of the last of the huge,
flowerless, coal swamp trees, flowers had evolved and spread. Crab
grass, philodendron, mother-in-law's tongue, spiderwort, Indian
corn, pumpkins, tulips, coconut palms, and willow trees are all rep-
resentative of flowering plants. Although the Amazon rain forest
contains a concatenation of flowering plants, as little as ten thousand
to only 2 percent of its present
years ago that rain forest extended
area. Like mammals, flowering plants-especially florid tropical
jungles-are recent evolutionary phenomena.
Some plants have evolved such a high degree of interdependency
with humans that they no longer survive in the wild. A graphic ex-
ample of such intimacy rs Zea mays, Indlan corn. Evolved from
 The Transmutation of Sunlight 209

teosinte, an obscure American grass, corn of many varieties now

towers, in season, over people around the world. Simply by choosing
seeds from the sweetest and most bountiful cobs year after year,
people have rendered Zea entirely dependent. lJnless removed and
individually husked and planted by human hands or agricultural ma-
chinery, corn cannot reproduce. Absent this assistance, the kernels,
trapped inside the fibrous covering, never sprout.
The "revolution"-the vast increase in population due to de-
velopment of agriculture and, subsequently, cities-is, from a bios-
pheric perspective, a major success story for flowering plants. Like
the Atta ants that tend their fungal gardens in rows, human inge-
nuiry and resources-farm animals, fossil-fuel-driven tractors, fer-
tilizers, irrigation, and biotechnological apparatus-have been de-
voted to maintaining the livelihood of our favorite plants. Our
primate brains, evolving in a world of flowering plants, are still de-
voted to preserving and extending that verdant, nurturing world.
Our attractions to angiosperms are deep and instinctive-so much
so that bottled essential oils are sold as perfumes, foods and drinks
are artificially fruit-flavored, and clothes and toys are dyed shades
of red, yellow, and orange-"warm" colors used in the original ad-
vertising campaign by which plants produced attractive and tasty in-
centives that enticed animals to do for them much of the work of
ferti-lization.
Plants, too, have animals to disperse their offspring. We eat grapes
but spit out the seeds, disseminating the plant. Bitter seeds and the
hard pits of fruits would merit the adjective "clever" were they the
fabrications of an organism with a hefry brain. Like colorful gro-
cery packaging, bright and flavorful fruits with inedible or discard-
able cores manipulate the animal into collecting and spreading the
offspring of the plant.
In an example of the growing intinracy among the many beings
cohabiting the biosphere, these immobile, muscleless, and brainless
beings-plants-have succeeded in appropriating the very powers
of restlessness and active perception that separate them from beings
 210 What ls Life?

to which they are commonly presumed to be inferior-animals. Like

the symbiotic anastomosis of branches on the tree of life, the merg-
ing of plant reproduction with animal sensitiviry and taste is a
demonstration of life's considerable powers of synergy and con-
vergence. Living beings not only compete and struggle; they also
associate and work together.

SOLAR ECONOMY
'We
bipedal mammals like to think ourselves king of the earthly hill,
the most evolved form of life. But the argument might just as well
be made in favor of flowering plants. They lack brains and speech-
but, then, they don't need them. They borrow ours.
With our vaunted intelligence we have been Johnny Appleseeds,
spreading fruit trees and grasses around Earth's surface. By tapping
more directly than any previous animal into past and present pho-
tosynthetic powers, we raise the stakes of life on Earth. For, make
no mistake about it, the solar economy has, with humans, entered
a new phase.
Peter Vitousek, using satellite imagery, estimates that 4o percent
of the ice-free land surface of the globe is under agricultural culti-
vation; very little arable land remains untilled (see plate z6). Hu-
manity annually uses the energy equivalent of r 8 trillion kilograms
of coal-about 3.6 metric tons for every man, wonlan, and child
on the planet. This total energy is used, in part, to retrieve 327,ooo
million kilograms of iron, 9o,ooo million kilograms of gypsum,
and sirnilarly staggering quantities of other materials. It is also used
to generate and retrieve an estimated 54o,ooo million kilograms of
wheat and 9z,ooo million kilograms of seafood.
As fossil fuels and solar energy integrate into factory and machine
production and into global husbandry and agriculture, more plants,
animals, and microbes come to depend on the technological system
now evolving. Nonrenewable resources are consunled, creating evo-
lutionary innovations in the form of new biospheric waste: insec-
 The Transmutation of Sunlight 2'l'l

ticides, polyvinylchloride, sryrofoam, rayon, and latex paints. The

gaseous by-products of burning long-buried energy sources perturb
or alter irreversibly the complex system of planetary physiology. Car-
bon dioxide accumulates in the atmosphere. Letting in visible light,
but trapping reflected heat, this greenhouse gas may increase plan-
etary temperatures-perhaps even melting polar ice, thus swamp-
ing coastal cities. Meanwhile, multiple extinctions follow frombazz-
sawing and bulldozing trees, killing some species directly but
upsetting far more by destructive incursion into their living space.
Nonetheless, the very energy our species uses to wreak habitat havoc
comes ultimately from photosynthesis. For good or evil, novelty or
status quo, nature is empowered by solar fire. The energy for vio-
lence also comes from plants.
Ever since Homo sapiens evolved, plants have fed, clothed, and shel-
tered us. From maternity ward vase to soft brown grave they ac-
company us on our biospheric journey. Borne on plants listing sun-
ward, flowers symbolize peace, life, beaury hope, femininiry and
the sun.
Flowers, like tropical fish in an aquarium, elevate and calm. Fo-
menting biophilia, they are mental medicine, provoking our spir-
its. But flowering plants-all plants-are more than decorative. They
are indispensable to an environment that can support humans. Their
descendants will keep our descendants company. Spider plants,
NASA reports, recycle trace pollutants in enclosed environments
such as a space capsule.'Water hyacinths and Nymphaea, awaterltly,
purifi, drinking water. Long trips into space are inconceivable with-
out plants to grow as food. Perhaps in seven generations your great,
great, great, great, great granddaughter will look down at her toes-
and see a wildflower poking up from a crack in the surface of Mars.

so, WHAT ls LIFE? Life is the transmutation of sunlight. It is the

energy and rnatter of the sun become the green fire of photosyn-
thesizing beings. It is the natural seductiveness of flowers. It is the
 212 | wn"t rs Life?
warmth of the tiger stalking the jungle in the dead of night. Green
fire converts wildly to the red and orange and yellow and purple
sexual fire of flowering plants. Expanding, developing lignin, green
beings raised up the biosphere and spread it horizontally. As fossils
these beings trapped the original gold of the sun, stocking wealth
oniy recently released in the human crucible of the soler economy.
But the arrow in all these transformations must eventually become
a loop that encloses the autopoietic exigencies of plants. We may
be an intelligent life form but our very intelligence depends on that
extension of ourselves that we now nurture as photosynthetic al-
lies. The prodigious grasslands, the lofry forests, and the lush gar-
dens are no mere backdrop to our machinate cleverness. Rather, the
plants provide the sustenance and energy upon which our apish
species unalterably depends. As life transmutes solar fre into all the
material and energetic cycles of the biosphere, we pay homage to
the ingenious ascension of the living plant.
 SENTIENT SYMPHONY

Owing to the imperfection of language, the offspring is termed

a new animal; but is, in truth, a branch or elongation of the
parent, since a part of the embryon animal is or was a part of
the parent, and, therefore, in strict language, cannot be said to
be entirely new at the time of its production, and, therefore, it
may retain some of the habits of the parent system.
ERASMUS DARWIN

They say that habit is second nature. Who knows but nature
is only first habit?
BLAISE PASCAL

Thinking and being are one and the same.

PARME N IDES

A DOUBTE LIFE

What Two crucial traits are that liG produces (autopoietically

is life?
self-maintains) and reproduces itself.Then there is inherited change:
DNA and chromosome mutation, symbiosis, and sexual fusion of
growing life when combined with natural selection means evolu-
tionary change. Nonetheless, autopoiesis, reproduction, and evolu-
tion only begin to encompass the fullness of life.
We have glimpsed ways of describing what life is: a material
process that sifis and surfs over matter like a strange, slow wave; a
planetary exuberance; a solar phenomenon-the astronomically lo-
cal transmutation of Earth's air, water, and received sunlight into

213
 214 What ls Life?

cells. Life can be seen as an intricate pattern of growth and death,

dispatch and retrenchment, transformation and decay. Connected
through Darwinian time to the first bacteria and through Vernad-
skian space to all citizens of the biosphere, life is a single, expand-
ing nerwork. LiG is matter gone wild, capable of choosing its own
direction in order to forestall indefinitely the inevitable moment of
thermodynamic equilibrium-death. Life is also a question the uni-
verse poses to itself in the form of a human being.
Life is manifest on Earth as five kingdoms, each revealing from a
different angle this mystery of mysteries. In a very real sense, life is
bacteria and their progeny. Every available piece of real estate on
this planet has become inhabited by subjects of the Kingdom Mo-
nera: by the enlightened producer, the tropical transformer, the po-
lar explorer. Life is also the strange new fruit of individuals evolved
by symbiosis. Different kinds of bacteria merged to make protoc-
tists. When conspecific protoctists merged the result was meiotic sex.
Programmed death evolved. Multicellular assemblages became an-
imal, plant, and fungal individuals. Life is thus not all divergence and
discord but also the coming together of disparate entities into new
beings.Nor did life stop at complex cells and multicellular beings.
It went on, forging societies and communities and the living bio-
sphere itself.
Life is moving, thinking matter, the power of expanding popula-
tions. It is the playfulness, precision, and wit of the animal kingdom-
which is a marvel of inventions for cooling and warming, moving
and holding firm, stalking and evading, wooing and deceiving. It
is awareness and responsiveness; it is consciousness and even self-
consciousness. Life, whose sinuous course was determined as much
by wily curiosiry as by historical contingency, is the flapping fin and
soaring wing of animal ingenuiry the avant-garde of the connected
biosphere epitornized by members of Kingdom Animalia.
Life is the transmigrator of matter, in which task fungi serve as
the closer of loops, making fungal food of plant and animal waste.
Life thus seeks out the underworld of soil and rot as much as the
sunny vistas enjoyed by photosynthetic beings. Life is a network of
 Sentient Symphony I ZIS

cross-kingdom alliances, of which Kingdom Mychota is a subtle,

seemingly crafty participant. It is an orgy of attractions, from the
deceptiveness of counterfeit fungal "flowers" to the delights of
trufHes and hallucinogens.
Life is the transmutation of energy and matter. Solar fire trans-
mutes into the green fire of photosynthetic beings. Green fire trans-
forms to the red and orange and yellow and purple sexual fire of
flowering plants, specialists in cross-kingdom persuasion. Fossilized
green fire is hoarded in the human cubicle of the solar economy.
Life is incessant heat-dissipating chemistry. And life is memory-
memory in action, as the chemical repetition of the past.
These halting descriptions approach but stop short of any final
definition of life. 'We will not proffer any last word, final judgment
because life will self-transcend; any definition In day-to-
slips away.
day adjustment and learning, in long-term action and evolution, in
interaction and coevolution, organic beings go beyond themselves
in the sense that they become more than what they were. Storing
and redistributing the energies of the sun, life displays ever greater
levels of activity and complexiry. Who can guess what liG might
make of itself if and when it expands to remake a greater part of
the universe into its home?
A1l organisms lead multiple lives. A bacterium attends to its own
needs in the muds of a salt marsh, but it is also shaping the envi-
ronment, altering the atmosphere. As communiry member it re-
moves one neighbor's waste and generates another's food. A fun-
gus goes about its business amid forest detritus, as it perforates the
leaf of a nearby tree and helps close the loop in the biospheric flow
of phosphorus. From one point of view we humans are ordinary
mammals; from another, a new planetary force.
Like other animals, we eat, urinate, defecate, copulate; Iike them,
we have descended from merged bacteria and meiotic ferrilizing pro-
toctists. Like other mamnealian species, Homo sapiens should expect
to endure for maybe another rr,vo million years or so-as the aver-
age species life span for mammals in the Cenozoic is less than three
million years. A11 species disappear: they extinguish or diverge to
 2'.t6 What ls Life?

form two or more descendant species. No animal species from Cam-

brian times is still alive today.
Perhaps Homo sapiens will diverge into rr,vo ollspring species differ-
ing as much from us today as we differ from chimps, Pan troglydites.
Such divergence may even be accelerated by technology. Human
descendants, their nervous systems incorporated into durable robotic
shells, may observe x-ray emissions of stars with telescope eyes as

they cling to interplanetary spacecraft. Perhaps some ex-humans

will free their genes of inherited disease or transcend-by genetic
manipulation-normal intelligence. Others, dwelling on planets
with higher or lower gravities, might undergo dramatic weight gains
or losses, with altered bone mass and respiratory systems and re-
arranged internal organs.
Many scenarios are imaginable. But, whatever we beconle, our
successors will retain traces of the past, which is our present. Con-
sider: even if some new biological weapon were able to instantly
vaportze all your animal cells, "you" would not disappear. As Clair
Folsome (r932-r988) mused:

What would remain would be a ghostly irrage, the skin outlined by

a shimmer of bacteria, fungi, round worms, pin worms, and various
other microbial inhabitants. The gut would appear as a densely packed
tube of anaerobic and aerobic bacteria, yeasts, and other microor-
ganisms. Could one look in more detail, viruses of hundreds of kinds
would be apparent throughout all tissues. 'We are far from unique. Any
animal or plant would prove to be a similar seething zoo of rnicrobes.l

We share more than 98 percent of our genes with chimpanzees,

sweat fluids reminiscent of seawater, and crave sugar that provided
ancestors with energy 3,ooo million years before the first space sta-
tion had evolved. 'We carry our past with us.
But now, aggregated into electronically wired cities, hun.rans have
begun retooling and transforming life on a planetary scale. Some
futurists claim that we have become exempt from mere animal evo-
lution. Are we not more than pretentious apes in fancy clothes?
 sentient Symphony I Xl
Do we not have music, language, culture, science, computer tech-
nology?
Rebuffed by Katharine Hepburn for a gin-drinking binge in the
film The Afrkan Queen, Humphrey Bogart abdicates responsibiliry
for his gruffbehavior. 'A man takes a drop too much once in a while,"
he explains, "it's only human nature."
" I',lature," replies Hepburn primly, looking up from her Bible, "is
what God put us on this Earth to rise above."
Self-transcending life never obliterates its past: humans are ani-
mals are microbes are chemicals. The view that we are "more" than
animals does not contradict the materialistic perspective underpin-
ning science. Life is less mechanistic than we have been taught to
believe; yet, since it disobeys no chemical or physical law, it is not
vitalistic. While we sense in ourselves a great degree of freedom,
all other beings, including bacteria, also make choices with envi-
ronmental consequences. Stored and transformed in life, the en-
ergy of sunlight powers cell growth, sex, and reproduction of higtrly
similar life forms. All living beings may share our own feeling of
free will.
Life on Earth is a complex, photosynthetically based, chemical
system ftactally arranged into individuals at different levels of or-
ganization. 'We cannot rise above nature, for nature itself transcends.
Nature does not end with us, but moves inexorably on, beyond
societies of animals. Global markets andEarth-orbiting satellite com-
nrunication, wireless telephones, magnetic resonance imagery, com-
puter net\,vorks, cable television, and other technologies connect us.
Indeed, people already form a more-than-human being: an inter-
dependent, technologically interfaced superhumaniry. Our activi-
ties are leading us toward something as far beyond individual people
as each of us is beyond our component cells.
Now, at the end of this century's hot and cold wars, we com-
municate freely across national boundaries at the speed of light, via
telephone and computer. News flashes around the world. But these
social changes at the start of the new millennium pale beside the
 214 What ls Life?

sweeping biological changes. The Phanerozoic eon, which began

more than 5oo million years ago with widespread predation and de-
fense against that predation in the form of animals with shells, is
ending. Evolutionary movements that made eukaryotes out of bac-
teria and animals from protoctists are now repeating on a planetary
scale. Humaniry is transforming from a sociery into a new level of
organic being. Our populations are beginning to behave as the brain
or neural tissue of a global being. As we become more populous
and sedentary, our human and technology-extended intelligence be-
comes part of planetary life as a whole.
The facts of life, the stories of evolution, have the power to unite
all peoples. By integrating the data of thousands of scientists, and
by cultivating the doubt and skepticism that is the epitome of sci-
entific inquiry, the cultural invention called science could provide
a more compelling, if ever corrigible, description of the world than

do parochial myths and divisive, faith-demanding religious traditions.

This hardly means that scientists are always correct. Yet the most
meaningful story of existence for future humaniry is more likely to
come from the evolutionary worldview of science than from Hin-
duism, Buddhism, Judeo-Christianiry or Islam. The dual under-
standing of scientific inquiry and creation myth could become a sin-
gle view: a science tale rich both in verifiable fact and personal
meaning.

cHotcE

In a truly evolutionary psychology, spirit and mind are not celestial

sprinklings but sovereign to living matter. Thought derives from no
world but this one; it comes from the activiry of cells.
When offered a variery of foodstuffi, swimming bacteria, ciliates,
mastigotes, and other mobile microbes make selections-they
choose. Squirming forward on retractable pseudopo ds, Amoeba pro-
teus frnds Tbtrahymena delectable but avoids Copromonas. Paramecium
prefers to gobble small ciliates, but if starved for these and other
protists it reluctantly feeds on aeromonad and other bacteria.
 Sentient Symphony 219

Although "merely" protoctists, foraminifera (one of the most di-

verse groups of fossil-forming organisms) make an astounding va-
riery of magnificent shells. Foraminifera without their shells resemble
amebas with very long, thin pseudopods. The shells are formed from
sand, chalk, sponge spicules, even other foram shells. To appropri-
ate their cell-shell homes, some forams agglute whatever particles
are available from the environment togetherwith an organic cement.
Observations reveal, however, that, when presented with a hodge-
podge of different particles, foraminifera make distinct choices based
on shape and size-Spiculosiphon, for example, passes over much of
the motley sediment, selecting only sponge spicules to make its test
or shell.z Without brains or hands, these determined protists choose
their building materials.
Smaller still, chemotactic bacteriajust t\,vo microns long can sense
chemical differences. They swim toward sugar and away ftom acid.
A chemotactic bacterium can smell a difference in chemical con-
centration that is a mere one part in ten thousand more concen-
trated at one end of its body than at the other.
Biochemist Daniel Koshland explains the spiritual leanings of
prokaryotes:

"Choice," "discriminarion," "memory," "learning," "insrinct,"'Judg-

ment," and "adaptation" are words we normally identifr with higher
neural processes. Yet, in a sense, a bacterium can be said to have each
of these properties. . . . it would be unwise to conclude that the analo-
gies are only semantic since there seem to be underlying relationships
in molecular mechanism and biological function. For example, learn-
ing in higher species involves long-term events and complex interac-
rions, but certainJy induced enzl,rne formation must be considered one
of the more likely molecular devices for fi-xing some neuronal con-
nections and eliminating others. The difference berween insrinct and
learning then becomes a matter of rime scale, not of principle.3

Microbes sense and avoid heat, move toward or away from light.
Some bacteria even detect magnetic fields. They harbor magnets
aligned in a row in their tiny, rod-shaped bodies (fig. t 8). That bac-
teria are simply machines, with no sensation or consciousness, seems
 220 What ls Life?

FIGURE 18. Magnetotactic bacterium remnant showing internal magnetosomes

(photo taken with an electron microscope). These cells, able to orient themselves mag-
netically to the north or south pole, exemplify the sensitivity of living substance at all
levels, scales, and kingdoms. Perception, choice, and sensation apply notjustto hu-
man beings or animals but, if they apply at all, they apply to all life on Earth.

no more likely than Descartes's claim that dogs suffer no pain. That
bacteria sense and act, but with no feeling, is possible-but ulti-
mately solipsistic. (Solipsism is the idea that everything in the world,
including other people, is the projection of one's own imagination.)
Cells, alive, probably do Gel. Indigestible mold spores and certain
bacteria are rejected by protists. Others are greedily gobbled. At even
the most primordial level living seems to entail sensation, choosing,
mind.
Darwin formally distinguished "natural selection" (referring to
interactions between nonhuman life and its environment) from
human-generated "artificial selection" (the aesthetic or functional
choices of pigeon fanciers, dog breeders, and agriculturalists). But
"natural" selection is, in a way, more "artificial," and far less me-
 Sentient Symphony I ZZI
chanical, than Darwin implied. The environment is not inert. Self-
awareness isnot confined to the space berween human ears. Non-
human beings choose, and all beings influence the lives of others.
Humans, we are told, are special. We have upright posture (a1-
lowing us to think of ourselves as literally "above" other species).
We have opposable thumbs (man the tool user), linguistic abilities
(man the storyteller), a superanimalistic soul (Descartes's distinction).
'We 'W'estern
have, at least in culture, a tradition of seeing ourselves
as being in a position of moral stewardship over the rest of life. Even
in the absence of God, we imagine ourselves to possess a unique
capacity to destroy the planet (via nuclear weaponry) or to swiftly
change atmosphere and climate.
Even such an ardent foe of the idea of progress in evolution as

Stephen Jay Gould (and he is not alone) proposes that whereas hu-
mans can evolve quickly through "cultural selection," all other forms
of life on Earth are shackled to the ancient, plodding system of "nat-
ural selection." But the sheer number of traits listed to explain hu-
man uniqueness is enough to arouse suspicion. Among the dazzling
array of reasons implying our superioriry over the rest of life, one
scientific argument stands out to us in curious contrast to the rest:
humans are the only beings capable of wholesale self-deception.
This claim is based on early humans' presumably delusionary be-
lief in the afterlife. Before written history our ancestors buried their
dead with food, weapons, and herbal medicines of litde use to corpses.
How ironic that we, in seeking examples of our superiority over the
rest of life, have finally congratulated ourselves on a trait that threat-
ens to negate all the others! Although members of other species trick
one another, humans are the expert self-deceivers: as the best sym-
bol users, the most intelligent species, and the only talkers, we are
the only beings accomplished enough to fully fool ourselves.

LITTLE PURPOSES

Freud's understanding of unconsciousness as repression-painful

memories are pushed away from the conscious mind-has diverted
 222 What ls Life?

attention from another way in which actions become unconscious.

Not avoidance but extended focus can make an action automatic,
second nature. A speech is "learned by heart." A practiced rypist no
longer glances at the keyboard. Virtually any activiry when memo-
rized subsides from conscious attention. The heart pumps, the kid-
neys filter in autonomic quasi perfection. Over breathing and swal-
lowing, normally automatic, the willful organism can exert some
voluntary control and modulation.
Now here is a strange thought: Perhaps we mammals remain un-
conscious of inborn physiology because, under pressure of survival,
our ancestors consciously practiced their skills to unconscious per-
fection. Although modern science does not yet offer us a mecha-
nism that transmits the learned habits of one generation to the phys-
iology of the next, experience shows that conscious can become
unconscious with repeated action. The gulf berween us and other
organic beings is a matter of degree, not of kind. Thken together,
the vast sentience comes from the piling up of little purposes, wants,
and goals of uncounted trillions of autopoietic predecessors who
exercised choices that influenced their evolution. If we grant our
ancestors even a tiny fraction of the free will, consciousness, and
culture we humans experience, the increase in complexiry onEarth
over the last several thousand rnillion years becomes easier to ex-
plain: life is the product not only of blind physical forces but also
of selection, in the sense that organisms choose. All autopoietic be-
ings have two lives, as the country song goes, the life we are given
and the life we make.
In the nineteenth century Samuel Butler (1835-r9oz), English
author, painter, and musician, challenged Charles Darwin's account
of evolution. Butler, who had many arguments with his father, left
for New Zealand to become a sheep farmer after completing his ed-
ucation at Shrewsbury and St. John's College, Cambridge. Excited
by Darwin's Origin o;f Species when he first read the book in New
Zealand, Butler gradually grew disenchanted with it. A scholarly
rebel who satirized sociery and explored the origins of religion, But-
 SentientSymphony I ZZI

ler accepted evolution but rejected Darwin's presentation. He be-

gan to suspect a dogmatism in the march of science as narrow as,
but more insidious than, that of the church. Reading previous evo-
lutionists, including Darwin's grandfather Erasmus, Butler accused
the younger Darwin of failing to acknowledge his intellectual
debts.
Darwin's schooling at Shrewsbury had been under the famous
headmaster Dr. Butler-Samuel Butler's grandfather. The younger
Butler claimed Darwin pretended to know little of his predeces-
sors, and in the first editions of Origin of Species made it seem, upon
returning from his famous Galapagos voyage, that the theory of nat-
ural selection had simply "occurred" to him. Butler pilloried Dar-
win's "Historical Sketch of the Progress of Opinion on the Ori-
gin of Species," which Darwin included in the second (r 86o), third
(r86r), and fourth (r866) editions of Origin. Darwin apologized
for this entry, described as "brief but imperfect." By the sixth and
final (t 872) edition the qualification was simply that it was "brief"-
carrying the implication that the historical sketch had perhaps grown
more nearly perfect in the interim. Butler disagreed.
What irked Butler most was Darwin's overly mechanical portrayal
of the evolutionary process. Darwin, Butler quipped, had "taken
the life out of biology." To be palatable to a religious populace in
the Victorian era Darwin's evolution needed a credible scientific
mechanism. Because the most respected achievement of the time
was the physical science of Isaac Newton, Darwin portrayed evo-
lution just as Newton had portrayed gravity: the result of abstract
principles and mechanical interactions.
Although best known as author of Erewhon and the posthumously
published The Way of All Flesh (an influential exploration of inter-
generational struggle), Butler Glt his greatest contribution was to
evolution theory. Retreating from Darwin's neo-Newtonian pres-
entation of organic beings as "things" acted on by "forces," Butler
presented sentient life as making numberless tiny decisions-and thus
responsible in part for its own evolution. Today Butler's view of the
 224 What ls Life?

sum ellbct of little purposes escapes reprobation only where hu-

-We
mans are concerned. consider ourselves forward-thinking cul-
tural beings, able to lay the flesh of the future upon the bones of
imagination. 'W'e even believe we can govern evolution. The rest of
life we dismiss as exempt from such Promethean foresight. Other
organic beings are portrayed as the result of physicochemical forces
or unmediated genetics-too inert, by insinuation, to play a for-
mative role in their own development. Butler begged to differ.
Butler's well-honed arguments and flashy polernics flouted the dry
scientific prose of his Victorian day. One Butlerian theme stands out:
living matter is mnemic, it remembers and embodies its own past.
Life, according to Butler, is endowed with consciousness. memory,
direction, goal-setting. In Butler's view all life, not just human IiG,
is teleological; that is, it strives. Butler claimed that Darwinians
missed the teleology, the goal-directedness of life acting for itself.
In throwing out the bathwater of divine purpose, Darwin discarded
the baby of living purposefulness.
No photosynthetic bacterium decided one day to become a wil-
low tree. Amoeba proteus does not today set out to make itself into
a mouse; it knows only that the swimming Tbtrahymena it relent-
lessly pursues is tasty. Ameba-knowledge at this level of sensing and
moving generates a million little such willful acts.These are sumcient
for evolution to work its wonders.
Life's purposes are grandiose only in the aggregate, and in retro-
spect. lJp close and confined in time they are ordinary. Nonethe-
less, no organic being is a billiard ball, acted upon only by external
forces. All are sentient, possessing the internal teleology of the au-
topoietic imperative. Each is capable, to varying degrees, of acting
on its own.

BUTLER'S BLASPHEMY

In the second of four books published at his own expense, Samuel

Butler discussed the evolutionary views of Erasmus Darwin, Jean-
 Sentient Symphony I ZZS

Baptiste Lamarck, Georges Louis Leclerc Buffon, and others. Eao-

lution, Old and New (published in r 879) was thus a fitting title. In a
circuitous series of newspaper letters and essays Butler published crit-
icism of Darwin's work: he accused the great man of evading the
credit owed Grandpa Erasmus and of mechanizing life. Butler even
questioned Darwin's honesry. Butler, trying to put life back into bi-
ology, hoped Darwin would respond to his r 879 book or to his ear-
lier (r877) LiJe and Habit.
Butler asked Darwin if his thought on evolution were an inspi-
ration without precedent that had appeared to him from on high.
'Was
itjust the result of contemplation upon a great number of facts?
Butler argued that the aura of poet-evolutionist Erasmus, coupled
with far-flung reading of evolutionary ideas, must have also con-
tributed to Darwin's intellectual development. Whether Darwin was
a subtle master of self-promotion or Butler a paranoiac of scholas-
tic patience rnay never be resolved. But Butler was predisposed, by
his own rebellious nature, combined with Darwin's growing status
as an intellectual icon, to feel disappointment in the great man. A
biographical book aboutErasmus Darwin had appeared. It was trans-
lated from German, and Darwin approved the translation as accu-
rate. Butler read it and was alarmed to find that the translation into
English of the early French evolutionist Lamarck used precisely the
same words that Butler had used to translate Lamarck in Butler's own
book Euolution, Old and New.There was also a comment that those
who wished to revive evolutionary thought prior to Charles Dar-
win showed "a weakness of thought and a mental anachronism
which no one can enr,ry'." Butler believed he was being attacked
obliquely in a fashion that precluded open debate; he believed that
Darwin himself might have been responsible for the dismissal of the
supposedly antiquated evolutionary ideas. He confronted Darwin
first by letter, then in the newspaper. In a letter to Butler, Darwin
explained that such alterations in translation as those made in the
book about his grandfather were so corunon that "it never occurred
to me to state that the article had been modified."
 226 What ls Life?

Darwin's family, and Thomas Huxley, urged Darwin not to re-

spond to any Butler criticism or personal letters. He didn't. Yet bi-
ographical records show that he drafted two responses, never sent.
In the second Darwin wrote he "could explain distinctly how the
accident arose, but the explanation does not seem to me worth giv-
ing. This omission, as I have already said, I much regret."a Fasci-
natingly, Darwin vacillates even here, berween the rwo modes of
explanation Butler was to neatly summarize in one of his books as
Luck, or Cunning: on the one hand, something was forgotten, there
was a lapse of memory; on the other hand, there was an "accident,"
something was "accidentally omitted." Even here, close up, we see
Darwin hedging, unable to characterize whether an event-and, in-
deed, one of his own doing-was the result of luck (chance, the
"accident") or cunning (choice and design), the very theory Butler
accused him of ignoring.
We agree with Butler that life is matter that chooses. Each living
being, Samuel Butler argued, responds sentiently to a changing en-
vironment and tries during its life to alter itself. But Living beings
cannot effect changes with great efficiency. No light bulb appeared
one day over the mammal that chose to become human. Rather,
gradually, in tiny increments, living systems with nonnegotiable
needs for food, water, and energy transformed themselves in wily
and persistent ways.
What theologians called design, and considered otherworldly, was
for Butler the result, in part, of Earth-bound thinking matter. The
analogy of the writer comes to mind: she is blocked, with only a
vague notion of what to write. Nonetheless, by following gram-
mar, spelling, and syntax, by adding word to word, something pur-
poseful emerges. The writerly result is not entirely her own because
she complies with the rules of language. Similarly, no life flouts any
law of physics, chemistry, or thermodynamics. As the decisions of
writers exist in the lexical world, so the choices of living beings ex-
ist in the material world. Neither are absolute, and yet the deeper
rules of matter, on the one hand, and language, on the other, im-
pose structures that permit overall designs to arise, not perfectly com-
 Sentient Symphony 227

plete but as accumulations of large numbers of minor, individual

decisions.
Butler believed minute changes effected by organisms on their
environment begin as conscious pursuits, but end as unconscious
practice. For Butler, amebas too have their little wants, their little
spheres of influence, their little "tool-boxes" with which they ma-
terially change their environments, pursue their little goals, and build
their little houses. This possibiliry is not ruled out by modern sci-
ence. Nobel Prize-winning Danish physicist Niels Bohr (r885-
ry62), in discussing the striking utilization by organisms of their
"past experience for reactions to future stimuli," contended that,
despite the success of physics-based mechanistic biology, there was
a need for description that includes "purposiveness."
'We
must realize that the description and comprehension of the closed
quantum phenomena exhibit no fearure indicating that an organiza-
tion of atoms is able to adapt itself to the surroundings in the way we
witness in the maintenance and evolution of living organisms. Fur-
thermore, it must be stressed that an account, exhaustive in the sense
of quantum physics, of all the continually exchanged atoms in the
organism not only is infeasible but would obviously require observa-
tional conditions incompatible with the display of life . . . it is evi-
dent that the attitudes termed mechanistic and finalistic [that is, pur-
poseful] do not present contradictory views on biological problems,
but rather stress the mutually exclusive character of observational con-
ditions equally indispensable in our search for an ever richer descrip-
tion of life.5

For Butler living matter can "memorize" its behavior not only
on the ontogenetic level of individual experience but also on the
phylogenetic level of species history. The transition between on-
togeny (the development of an individual life) and phylogeny (per-
sistence and change of many individuals through time) is relative.
The difference benveen the same individual newly born and at age
eighry is greater, Butler argued, than that between a newborn in-
fant of one generation and a newborn infant of the next.
Reptiles shed their skins; insects reshuffie their proteins in the pupa
 228 What ls Life?

stage. The corpse is replaced, with some overlap, by its grandson.

W'e moderns accept that a caterpillar "metamorphoses" into a but-
terfly, yet we employ the term "death" for what happens to grand-
father's body. Nevertheless, with both the metamorphosing insect
and the dying grandparent, the fresh body ofyouth reappears. Each
of us is entitled to think we die, suggests Butler, but the demarca-
tion is highly arbitrary: a parent contributing to the flesh of a child
is a prolongation, not an abrupt end, to biological continuiry. The
"individual" is not so complete, in time, as we have been taught.
Butler believed unconsciousness applies not only to adult human
beings but throughout many difrerent levels of living organization.
The most important tasks, the ones most often repeated, have be-
come the most unconscious, the most "physiological." The pump-
ing of blood is too ancient and important to be unlearned or
modified easily by mere lassitude or an act of will. Divorce from
playful consciousness insures that routine but critical activities are
competently performed. Treated as work, effected regularly and au-
tomatically, important physiology is not exposed to experimenta-
tion that might destroy it. The once-conscious process of steering
an automobile recedes to the unconscious while the mind attends
to other matters.
No concert pianist strives to hit the keys in order. The long pe-
riods of former conscious striving ingrain the knowledge into the
musician's fingers. Dancers call their talent "muscle memory."
Choice and practice become smooth habir.
Cells, with eons of practice, do not consciously decide to respire
oxygen or reproduce by rnitosis. They, or the bacterial remnants that
compose them, may once have so strived. The more recent the ad-
dition of a habit to life's physiological repertoire, rhe more likely it
is to be conscious or at least subject to conscious interference. No
longer do animals, plants, and fungi directly respond to the meta-
bolic pathways by which their cells obtain energy when oxygen gas
reacts with food hydrogen atoms to produce water and carbon diox-
ide. Such metabolic behavior is submerged in the permanent un-
 sentient Symphony I ZZI

conscious of modern organisms. Oxygen-hydrogen reactions per-

formed by mitochondria, which were once free-living bacteria, are
a chemical feat that has not ceased for z,ooo million years, ever since

oxygen respirers responded to an environment transformed by oxy-

genic cyanobacteria. By contrast, peristalsis in digestion-also an un-
conscious phenornenon but one of which mamrnals tend to be
aware-evolved in animal ancestors long after the microbial stage.
Swallowing, chewing, and speaking are behaviors that have been
learned far more recently, and in that order.
Butler's theory of unconscious memory holds that all beings are
capable of forming habits, some of which-upon much repetition-
'We
become physiologically ingrained over the course of evolution.
do not remember, writes Butler, when first we grew an eye. Some-
day, he muses, so many of us will have learned to read and write,
so often, that we will be born that way. Perhaps a future William
Harvey will be required to uncover the details of how the learning
of reading became a physiology, just as Harvey illuminated the cir-
culation of the blood. Here we disagree with Butler. It seems doubt-
ful that children will ever be born reading and writing. However,
if one substitutes television for writing one can see that Butler is al-
ready on target: children are born, more or less, watching televi-
sion; and television, put together from technological products made
by companies all over the planet, is already so complicated that few
humans understand it in detail.

HABITS AND MEMORY

Given free will and the status of living beings as open thermo-
dynamic systems, one should not be too quick to use classical physics
to justify an understanding of life as a mechanical phenomenon. A
general property distinguishing life from nonliving matter is its his-
torical coherence, including the potential to evolve. By exporting
disorder, randomness, and entropy to their surroundings, living sys-
tems increase local complexiry intelligence, and beaury building on
 230 What ls Life?

the past and planning for the future. Organisms that find new means
of extracting energy and matter for the perpetuation of their form
will tend to be preserved, leading to increasingly curious and cre-
ative beings.The hints and hunches must be replaced by firm detail,
yet the extraordinary storage and transmission processes of life for
molecular herediry as well as cultural information, may be robust
enough to encompass the phenomenon postulated by Butler: phy-
logenetic "memorization," the conversion of the conscious strivings
of one generation into the activities and, eventually, the physiologies
of the next.
Although we fail as yet to see how an organism's or even a species'
voluntary habits can become the physiology of a future generation
via a material basis of herediry we are fascinated by Butler's sug-
-We
gestion. know, for example, that many organic beings acquire
new heritable traits by symbiogenesis and that a vast array of oth-
ers, not only people, are capable of learning. Ecosystems grow in-
creasingly complex and sensitive; processes practiced in them re-
peatedly by one generation may become easier for the next. More
open-minded investigation is needed. Objections may be leveled
against Butler's ideas, yet he cannot be accused of the atavistic think-
ing which clings to humaniry's separate status. Covertly consider-
ing ourselves divine, under the scientific rubric of "cultural evolu-
tion," or by dint of that other desperate euphernism, our "big
brains," we are probably now more ecologically impoverished than
we would be if, a century ago, we had adopted Butler's notion of
all life as a conscious continuum.
Butler did not object to evolution but to the loss of the richness
of the earlier, more lively views, in which living beings themselves
were involved in natural selection:

According to Messrs. Darwin and'Wallace, we may have evolution,

but are on no account to have it as mainly due to intelligent effort,
guided by . . . sensations, perceptions, ideas. 'We are to set it down to
the shuffling of cards. . . . According to the older men, cards did in-
deed count for ntuch, but play counted for more. They denied the
 Sentient Symphony 231

teleology of the time-that is to say, the teleology that saw all adap-
tation to surroundings as part of a plan devised long ages since by a
quasi-anthropomorphic being who schemed. . . . This concePtion
they found repugnant alike to intelligence and conscience, but,
though they do not seem to have perceived it, they left the door open
for a design more true and more demonstrable than that which they
excluded. . . . They made the development of man from the amoeba
part and parcel of the story that may be read, though on an infinitely
smaller scale, in the development of our most powerful marine en-
gines from the common kettle, or of our finest microscopes from the
dew drop. The development of the steam-engine and the microscope
is due to intelligence and design, which did indeed utilize chance sug-
gestions, but which improved on these, and directed each steP of their
accumulation, though never foreseeing more than a step or wvo ahead,
and often not so much as this.6

EXISTEN CT'S CELEBRATION

For nineteenth-century Englishmen of science it was natural and

expedient to invoke Newtonian mechanics and conceive of life as
Newton's matter: blind bits predictably responding to forces and nat-
ural laws. Like some piece of well-made clockwork, the world was
donated or its mechanism manufactured by a transcendent god-a
god that then stood outside its creation.
This was the new view of evolution: God, if it existed, was New-
ton's God. Not an active interloper in human afhirs, it was the god
of the mathematicians, the geometer god, who made the laws and
then sat by and watched those laws play out. But an older view left
room for a kind of god, too-a more active god. This was the view
that Samuel Butler attempted to resuscitate-that life itself was god-
like. There was no grand design, but millions of little purposes, each
associatedwith a cell or organism in its habitat.
To the neo-Newtonians, the Darwinians, free will had been all
but banished from the universe because the universe was portrayed
as a mechanism and mechanisms do not have consciousness. For
 232 | wnut ts Life?

Descartes, God continued to have consciousness and people did

insofar as they were in touch with God. But when Darwin showed
through painstaking work that people too could be explained by
the mechanism of natural selection, consciousness suddenly be-
came redundant in the human world as well. Butler brought con-
sciousness back in by claiming that, together, so much free will,
so much behavior becoming habit, so much engagement of mat-
ter in the processes of life, so many decisions of where, how, and
with what or whom to live, had shaped life, over eons producing
visible organisms, including the colonies of cells called human.
Power and sentience propagate as organisnrs. Butler's god is im-
perfect, dispersed.
We find Butler's view-which rejects any single, universal
architect-appealing. Life is too shoddy a production, both physi-
cally and morally, to have been designed by a flawless Master. And
yet life is more impressive and less predictable than any "thing"
whose nature can be accounted for solely by "forces" acting deter-
ministically. The godlike qualities of liG on Earth include neither
omniscience nor omnipotence, although an argument could be made
for earthly omnipresence.
Life, in the form of myriad cells, from luminescent bacterium to
lily-hopping frog, is virtually everl-'uvhere on the third planet. All
life is connected through Darwinian time and Vernadskian space.
Evolution places us all in the stark but fascinating context of the
cosmos. Although something odd nray lurk behind and before this
cosmos, its existence is impossible to prove. The cosmos, more daz-
zlingthan any sect's god, is enough. Life is existence's celebration.
Butler's forgotten theory intrigues us. The mind and the body are
not separate but part of the unified process of life. Life, sensitive from
the onset, is capable of thinking. The "thoughts," both vague and
clear, are physical, in our bodies' cells and those of other animals.
In comprehending these sentences, certain ink squiggles trigger
associations, the electrochemical connections of the brain cells. Glu-
cose is chemrcally altered by reaction of its components with oxy-
 sentientSymphony I ZAI

gen, and its breakdown products, water and carbon dioxide, enter
tiny blood vessels. Sodium and calcium ions, pumped out, traflic
across a neuron's membranes. As you remember, nerve cells bolster
their connections, new cell adhesion proteins form, and heat dissi-
pates. Thought, like life, is matter and energy in flux; the body is its
"other side." Thinking and being are the same thing.
If one accepts this fundamental continuity berween body and
mind, thought loses any essential difference from other physiology
and behavior. Thinking, like excreting and ingesting, results from
lively interactions of a being's chemistry. Organism thinking is an
emergent properfy of cell hunger, movement, growth, association,
programmed death, and satisfaction. Restrained but healthy former
microbes find alliances to make and behaviors to practice. If what
is called thinking results from such cell interactions, then perhaps
communicating organisms, each of themselves thinking, can lead
to a process greater than individual thought. This may have been
what Vladimir Vernadsky meant by the noosphere.
Gerald Edelman and William Calvin, both neuroscientists, have
each proffered a kind of "neural Darwinism." Our brains, they say,
become minds as they develop by rules of natural selection.T That
idea may provide a physiological basis for Butler's insights. In the
developing brain of a mamrnalian fetus, some I o12 neurons each be-
come connected with one another in
roa ways. These cell-to-cell
adhesions at the surface membranes of nerve cells are called synap-
tic densities. As brains mature, over 90 percent of the cells die! By
programmed death and predictable protein synthesis, connections
selectively atrophy or hypertrophy. Neural selection against possi-
bilities, always dynamic, leads to choice and learning, as remaining
neuron interactions strengthen. Cell adhesion molecules synthesize
and some new synaptic densities form and strengthen as nerve cells
selectively adhere and as practice turns to habit. Selection is against
most nerve cells and their connections but it is nevertheless for a
precious few of them. Of course, more work is needed before the
physical basis of thought and imagination can be understood, but
 234 ] wf'rt ls Life?

selective death in a vast field of proliferating biochemical possibili-

ties may apply to rninds as it does to evolution.
The peculiar\ curved early embryos of birds, alligators, pigs, and
humans are remarkably similar. Developing from a fertile egg, all
display a stage with gill slits-whether the hatched or born animal
breathes oxygen from air or water. The slits that close behind the
ears in the human fetus attest to our corunon ancestry with fish,
whose gill slits function in the adult. Human embryos have tails.
Living matter "remembers" and repeats its origins to arrive in the
present. In a Butlerian world, the materials of living beings are
molded by liG, over and over, for millions of generations. Creating
a sense of dtjn uu, the embryo represents a once-unconscious process,

now again-at a different level-brought to consciousness.

SUPERHUMANITY

A transhuman being, superhumaniry is appearing, becoming part

of the sentient symphony. It is composed not only of people but of
material transport systems, energy transport systems, information
transport systems, global markets, scientific instruments. Superhu-
maniry ingests not only food but also coal, oil, iron, silicon.
The global network that builds and maintains cities, roadways,
and fiber-optic cables grows by leaps and bounds. In Nigeria, for
example, the population is expected to reach zr6 million by the
year 20ro, double that of 1988. Unchecked, such growth would
bring the number of Nigerians to more than ro,ooo million by the
ye^r 2rro-twice the present global human population. Our stu-
pendous population taps a significant proportion of the solar en-
ergy reaching Earth's surface. The raw energy of photosynthesis,
past and present, and transformed into edible plants, animal fod-
der, geological reserves, and human muscle and brain, supports the
of the transcontinental urban ecosystem and
massive construction
even-"biting the hand that feeds it"-the raztng of forests cap-
turing and converting solar energies. As the system expands using
 Sentient Symphony I ZIS

genetic and atomic technology, its operations become more elegant

and cohesive. The potential for disaster also increases.
Superhumanity is neither a simple collection of humans nor some-
thing other than aggregated humans and their devices. Plumbing,
tunnels, water pipes, electric wires, vents, gas, air conditioning ducts,
elevator shafts, telephone wires, fiber-optic cables, and other links
enclose humans in a rapidly growing net. The way superhumaniry
behaves is in part the result of uncountable and unaccountable eco-
nomic decisions made by people-singly and in groups-within
the context of an increasingly planetary capitalism. "The problem
with money," says a character in a recent film, "is that it makes us
do things we don't want to do."
Whether or not superhumaniry's tendencies are conscious beyond
us, individual humans should not be surprised if the aggregate of
planetary humaniry shows unexpected, emergent, seemingly pur-
poseful behaviors. If brainless bacteria merged into fused protists,
which cloned and changed themselves over evolutionary time into
civilization, what spectacle will emerge from human beings in global
aggregation? To deny the existence of superhumaniry by insisting
it is merely the sum of human actions is like claiming that a person
is merely the sum of the microbes and cells that constitute the body.8

EXPANDING LIFE

Life today is an autopoietic, photosynthetic phenomenon, plane-

tary in scale. A chemical transmutation of sunlight, it exuberantly
tries to spread, to outgrow itself. Yet by reproducing, it maintains
itself and its past even as it grows. Life transforms to meet the con-
tingencies of its changing environment and in doing so changes that
environment. By degrees the environment becomes absorbed into
the processes of life, becomes less a static, inanimate backdrop and
more and more like a house, nest, or shell-that is, an involved, con-
structed part of an organic being. The members of thirry million
species interacting at Earth's surface continue to change the world.
 236 What ls Life?

Coming to understand iife afresh, we find that species of organ-

isms diverge into new kinds, yet earlier patterns never entirely disap-
pear. Old life form.s, the bacteria that run the planet's ecology, are
supplemented but not replaced. Although every distinct variety of nu-
cleatedbeing-every species of plant, animal, protoctist, andfungus-
perishes, sirnilar new taxa evolve &om them or from their kind. Mean-
while, the underlying bacteria march symbiogenetically on.
We find that nature is not always "cutthroat," or, as the poet Al-
fred Tennyson put it, "red in tooth and claw." Living beings are
amoral and opportunistic, as befits their needs for water, carbon, hy-
drogen, and the rest. They are fractally repeating structures of mat-
ter, energy, and information, with a very long history. But they are
no more inherently bloodthirsry competitive, and carnivorous than
they are peaceful, cooperative, and languid. Lord Tennyson might
just as well have cast nature as "green in stem and leaf."
Among the most successful-that is, 2fg16l2n6-living beings on
the planet are ones that have teamed up. Moving inside (or perhaps
forcibly dragged inside) another cell, the cyanobacteria that became
chloroplasts in protoctist and plant cells weren't lost; they were trans-
forrned. So too with the mitochondria-once respiring bacteria-
that give your finger muscles the energy to turn this page. Former
bacteria, as themselves or parts of larger cells, are still the most abun-
dant forms of life on the planet.
The strength of symbiosis as an evolutionary force undermines
the prevalent notion of individualify as something fixed, something
secure and sacred. A human being in particular is not single, but a
composite. Each of us provides a fine environment for bacteria, fungi,
roundworms, rnites, and others that live in and on us. Our gut is
packed with enteric bacteria and yeasts that manufacture vitamins
for us and help metabolize our food. The pushy microbes of our
gums resemble department store customers before a holiday. Our
mitochondria-laden cells evolved from a merger of fermenting and
respiring bacteria. Perhaps spirochetes, symbiotically faded to the
edge of detectabiliry continue to squirm as the undulipodia of our
 Sentient Symphony I ZZZ

fallopian tubes or sperm tails. Their remnants may move in subtle

ways as our microtubule-packed brains grow. "Our" bodies are ac-
tuallyjoint properry of the descendants of diverse ancestors.
Individualiry is not stuck at any one level, be it that of our own
species or pondwater Amoeba proleas. Most of our dry weight is bac-
teria, yet as citizens swarming in crowded streets and office build-
ings, viewing television, traveling in cars, and communicating by
cellular and facsimile phone, humans disappear in a global swirl of
activiry overwhelmed by emergent structures and abilities that could
never be accomplished by individuals or even tribes of human pred-
ecessors. No single human can speak to another human, in real time,
thousands of miles away. No single human could stand on the moon.
These are emergent abilities of superhumaniry. Our global activi-
ties bring to mind the social insects, except that our "hive" is nearly
the entire biosphere.
Inextricably embedded in the biosphere, this superhuman sociery
is not independent. At its greatest extent life on Earth-fauna, flora,
and microbiota-is a single, gas-entrenched, ocean-connected plan-
etary system, the largest organic being in the solar system. The up-
per mantle, crust, hydrosphere, and atmosphere of Earth remain in
an organized state very different from that on the surfaces of our
neighboring planets. Photosynthesis, respiration, fermentation, bio-
mineralization, population expansion, seed germination, stampedes,
bird migration, mining, transportation, and industry move and al-
ter matter on a global scale. Life dramatically impacts the environ-
ment by producing and storing skeletons and shells of calcium phos-
phate and carbonate, by caching plant remains as coal and algae
residues as oil. Great layers of rninerals-sulfides of iron, lead, zinc,
silver, and gold-remain in place where they were precipitated by
hydrogen gas-producing bacteria.
Minerals not normally associated with life-aragonite, barite, cal-
fluorite-are produced as crystals and skeletons in-
cite, francolite,
side, and exoskeletons or shells outside, living organisms. Plants and
microbes induce the formation of "inanimate" substances such as
 238 What ls Life?

barite, iron oxides, galena (lead sulfide), and pyrite (iron sulfide, or
fool's gold). Humanity's cultures are ranked from a stone age
through an iron to a bronze age. Some argue that with the advent
of computers Earth has entered a "silicon age." But metallurgy pre-
ceded us: human metalworking followed the bacterial use of mag-
netite for internal compasses by 3,ooo million years. Pedomicrobium,
a soil bacterium found fossilized in gold samples, is thought to pre-
cipitate gold ions and thus accumulate gold particles in its sheath.
Next to hundreds of cubic kilometers of tropical reefs built by coral
and entire cliffs of chalk precipitated by foraminifera and cocco-
lithophorids, human technology does not seem uniquely grand.
Our destiny is joined to that of other species. When our lives
touch those of difrerent kingdoms-flowering and fruiting plants.
recycling and sometimes hallucinogenic fungi, livestock and pet
animals, healthful and weather-changing microbes-we most feel
what it means to be alive. Survival seems always to require more
networking, more interaction with members of other species, which
integrates us further into global physiology. Despite the apocalyp-
tic tone of some environmentalists, our species is on its way to be-
coming better integrated into global functioning. Though tech-
nology can poison humans and other organisms and stultify their
growth, it also has the capaciry to usher in the next major change
in biospheric organization.
Teamwork enabled life to spread on Earth: anaerobic microbes
joined to make the swimrning ancestors of protist colonists, masti-
gotes ingested but did not digest the mitochondria that allowed them
to invade oxygen-rich niches atEarth's surface, fungi and algae corn-
bined into lichens that colonized bare rock of dry land. The trans-
port of life to new planetary bodies will also require teamwork. As-
tronautics, computers, genetics, biospherics, telecommunicarions, and
other forms of human-sired technology will have to combine with
the predecessor technology-most significantly photosynthesis-
of other planetmates. The ultimate explosion of life onto its next
frontier-that of space-will rely on the new technology of life it-
 Sentient Symphony 239

self. Vivification and terraformation-the coming to life of other

planetary bodies-are not only human processes. Someday recycling
ecosystems inside spacecraft may feed humans voyaging to other
planets. If
humans are to reside in space or voyage beyond Earth's
orbit, the plants that feed them, the bacteria that digest for them,
the fungi and other microbes that recycle their wastes, and the tech-
nology that supports them will surely be along for the ride. The ex-
tension of our local thermodynamic disequilibrium into space nec-
essarily will involve representatives of all five kingdoms that have
forged new ecosystems, able to transfer energy and cycle matter in
isolation from the mother planet, the original functioning biosphere
of Earth.
The distinction between space colonization by machines alone or
by life with machines mirrors a New Zealand newspaper debate
Samuel Butler had with himself. Beginning in r86z Butler con-
tributed an anonymous article to The Press of Christchurch, New
Zealand. At the time, he was sheep farming in the Upper Rangitata
district of Canterbury Province. Entitled "Darwin on the Origin of
Species, a Dialogue," the unsigned article generated spirited protest.
Butlerjoined in, criticizing himself as well as others. Signing his di-
vided opinions under different names, Butler ultimately argued for
rwo diametrically opposed interpretations of machines.
In "Darwin among the Machines," a letter signed by one Cellar-
ius and appearing tn The Press on r 3 June r 863 , Butler held that ma-
chines were the latest form of life on Earth, poised to take over and
enslave their human masters; the rate of evolution and reproduction
of machines was prodigious, and without "war to the death" at once
it would be too late to resist their world dorninion. Then, in a z9
July r 865 article entitled "Lucubratio Ebria," Butler countered Cel-
larius by saying that human beings were not even human without
clothes, tools, and other mechanical accessories. Machines were not
a threat to human life, but its indissociable natural extension.e
If space vehicles do cut free of human influence, voyaging starry
skies as they reproduce on their own, then Cellarius and otherLud-
 24O What ls Life?

dites will be vindicated. If, however, machines in space flourish

not alone but as intelligent enclosures for a wide variety of other
life forms, then the author of "Lucubratio Ebria" will be proven
correct.
We place our bets on the latter. Machines, we believe, will flour-
ish in a tighdy meshed interface with life-not only human life but
a rich assortment of starlight-using life forms. People are essential
for making the export of life into the night fantastic even possible.
But, like the sperm tail which breaks off once the genetic message
enters the egg, so human beings are ultimately expendable. Even
without us, a hundred rnillion more years of sun-driven planetary
exuberance should be enough to get liG offthe Earth, star-bound.
Other technological species might evolve. Besides, not only humans
have begun space exploration.
Setting foot on the moon, Neil Armstrong proclaimed, "One
small step for man, one giant leap for mankind." True, in a sense;
but he overlooked vast numbers of bacteria on his skin and in his
intestine that stepped with him. Life has been expansionist from the
beginning. Once it gets a firm toehold in space it may kick off its
human shoes and run wild.

RHYTHMS AND CYCLES

We and many other animals sleep and wake in cycles that repeat
every twenfy-four hours. Some ocean protists, dinomastigotes, lu-
minesce when dusk comes, ceasing two hours later. So hooked are
they into the cosmic rhythm of Earth that even back in the labora-
tory, away from the sea, they know the sun has set. Many similar
examples abound because living matter is not an island but part of
the cosmic matter around it, dancing to the beat of the universe.
Life is a material phenomenon so finely tuned and nuanced to its
cosnric domicile that the relatively minor shift of angle and tem-
perature change as the tilted Earth moves in its course around the
sun is enough to alter life's mood, to bring on or silence the song
 SentientSymphony I Zn't

of bird, bullfrog, cricket, and cicada. But the steady background beat
of Earth turning and orbiting in its cosmic environment provides
more than a metronome for daily and seasonal lives. Larger rhythms,
more difiicult to discern, can also be heard.
Many rypes of life form encapsulating structures that protect them
from temporary dangers of the environment. Propagules of a wide
variety of rypes are miniaturized, viable representatives of mature
organic beings. They range from bacterial and fungal spores to pro-
toctist cysts; from plant spores, pollen, seeds, and fruit to the dry
eggs of some crustaceans, insects, and reptiles. As such propagules
proliferate, natural selection deals with them severely: many die or
fail to grow
Desiccation- and radiation-resistant, most propagules metabolize
at an exceedingly slow rate. Spores of bacteria may lay in wait for
a hundred yex15-11111i1 rain comes, or phosphorus abounds, or con-
ditions otherwise become less dry and more permissive. Without
any dormant seeds or resistant spores, humans survive extraordinary
environmental hazards. Houses, clothes, railroads, and automobiles
have made possible our expansion from the subtropical home to
colder climes. Analogous to spores. cysts, and seeds, these structures
protect us from harsh conditions.
Recycling greenhouses are enclosed dwellings that contain repre-
sentative collections of Earth's life. They detoxifi, poisons and trans-
form wastes into food and back again. One, designed by Santiago
Calatrava, will span the entire roof of the Cathedral of Saint John
the Divine in NewYork City. Such "artificial biospheres" miniatur-
ize crucial processes of the autopoiesis of the global ecosystem.
The global ecosystem is not an ordinary organic being. The global
system, like all living beings, is energetically open: solar radiation
comes steadily in, dissipated heat moves steadily out. But unlike other
beings, the global system is closed to material exchange. Apart from
the occasional incoming meteor or comet, nothing enters. Apart
from the occasionally stalled geologic churning here and there of
sediments into new crust and cooked gases, nothing leaves. A11 the
 242 What Is Life?

matter used by life is recycled matter-reappearing matter that is

never consumed.
No living cell or organism feeds on its own waste. Thus artificial
ecosystems have biological importance that exceeds architecture or
other human concerns. For the first time in evolutionary history,
the biosphere has reproduced, or better put, has begun reproduc-
ing itself through humankind and technology. The generation of
new "buds," materially closed living systems, within the "mother
biosphere" resembles the structure of a fractal.
From a "green" or "deep ecology" perspective, humans do not
dominate but are deeply embedded within nature. Artificial bio-
spheres are the first buds of a planetary organic being, as "man-
made" biospheres have the potential to duplicate biospheric, light-
dependent self-sufiiciency. People in NASA, the European Space
Agencies, governments, private industry, academia and elsewhere
are pondering these desiccation-resistant structures that sequester,
like new Noah's arks, samples of life-not in museums but in a liv-
ing, selGsustaining form. The largest of the recycling structures is
Biosphere z in the Sonoran desert at Oracle, Arizona. Ultimately,
closed ecosystems are not artiftcial at all, but part of the natural
processes of self-maintenance, reproduction, and evolution in a heat-
dissipating universe.
For an organic being to survive in space, food must be replen-
ished and waste disposal systems must work. Photosynthesis has
stored solar energy in rocks as reserves of kerogen, oil, gas, iron
sulfide, coal, and other substances. The planet's prodigal species now
expends these reserves, using the energy to spread its populations.
Homo sapiens spends the riches of eons; meanwhile, the rhythms of
Earth, building up and breaking down for ages, crescendo. Our cre-
ative destruction accelerates. But nature has not ended, nor does the
planet require saving. The technological dissonance marks no end
but a 1u11, a gathering of forces.
Global life is a system richer than any of its components.'We alone
among the animals build large telescopes and mine Archean dia-
 Sentient Symphony I Znl

monds. But, while our position cannot soon or easily be replaced,

we are not in charge. The diamonds are made of carbon, a main el-
ement of life since its inception some 4,ooo rnillion years ago; and
telescopes are lenses, parts of the compound eye of a metahuman
being that is itself an organ of the biosphere.
The continuous metamorphosis of the planet is the cumulative
result of its multifarious beings. Humankind does not conduct the
sentient symphony: with or without us, life will go on. But behind
the disconcerting tumult of the present nlovement one can hear,
like medieval troubadours climbing a distant hill, a new pastorale.
The melody promises a second nature of technology and life, to-
gether spreadingEarth's multispecies propagules to other planets and
the stars beyond. From a green perspective a keen interest in high
tech and the altered global environment makes perfect sense. It is
high noon for humanity. Earth is going to seed.
 Epilogue

There is not so much Life as talk of Life, as a general thing. Had

we the first intimation of the Definition of Life, the calmest of us
would be Lunaticsl
EMILY DICKINSON

Half a century ago the preeminent physicistErwin Schrcidinger, a

humane and thoughtful scholar, approached, as science, the ques-
tion of what life is. Working prior to the discovery of DNA and
before knowledge accumulated on how enzyme proteins and chem-
ical movement become the metabolism of life, Schrcidinger nonethe-
less inspired the very search that led to a materialistic explanation
'We
of living processes. are fortunate to engage Schrodinger's tradi-
tion after fifry additional years of scientific inquiry.
To function, the biosphere requires microbial diversiry; to feel
whole and at home most of us crave nature's variery. Perhaps we now
are more alarmed about the human prospect than was Schrcidinger
because we live on a more populated Earth. Humans today clearly
are threatened by the extinction, even before science can describe
them, of so many of our planetmates. Plastics spread, tropical rain
forests die out, coral reefs collapse.'We wonder whether the grow-
ing understanding of life's autopoietic tendencies for expansion
and control and the evolutionary history of planetary change in
the wake of rapidly spreading life forms will make individuals less
likely to buy packaged plastic goods, travel using fossil fuels, eat

245
 246 Epilogue

meat, or engage in other environmentally destructive activities. We

doubt it!
Knowledge about the varieties of life onEarth-life which, from
pond scum to tigress, is connected to us through time and space-
serves to inspire. That excess is natural but dangerous we learn from
the photosynthetic ancestors of plants. That movement and sensa-
tion are thrilling we experience as animals. That water means life
and its lack spells tragedy we garner from fungi. That genes are
pooled we learn from bacteria. Modern versions of our ancient
aquatic ancestors, the protoctists, display versions of the urge to cou-
ple, and of our capacity to make choices. Humans are not special
and independent but part of a continuum of life encircling and em-
bracing the globe.
Homo sapiens tends to dissipate heat and accelerate organization.
Like all other life forms, our kind cannot continue to expand lim-
itlessly. Nor can we continue to destroy the other beings upon whom
we ultimately depend. 'We must begin really to listen to the rest of
life. As just one melody in the living opera we are repetitious and
persistent. We may think ourselves creative and original but in those
talents we are not alone. Admit it or not, we are only a single theme
of the orchestrated life-form. 'W'ith its glorious nonhuman past and
its uncertain but provocative future, this life, our life, is embedded
now as it always has been, in the rest of Earth's sentient symphony.
Now, as before, life is empowered by the sun. It is a phenomenon
not only molecular but astronomic. Life is open to the universe and
to itself. In the tradition of Charles Darwin, Samuel Butler, Vladimir
Vernadsky, and Erwin Schrodinger, we can ask with curiosiry but
can answer only tentatively and with humiliry the question of what
life is, hoping, with you, that the search continues.
 Notes

CHAPTER 1. LIFE: THE ETERNAL ENIGMA

The epigraphs are from James E. Lovelock, The Ages of Gaia (New York:
W.'W. Norton, 1988), p. 16; and Robert Morison, "Death: Process or
Event," Science, zo August tg7o, pp. 694-698.
r. Erwin Schrcidinger, My View of the Woild (Cambridge: Cambridge
UniversiryPress, r967), p. 5.
z. Thomas Mann, cited in Frederick Turner, "Biology and Beaury," in
lncorporations, ed. Jonathan Crary and Sanford Kwinter (New York: Zone
Books, ry92), p. 4o6.
3. Arthur Koestler, Janus: A Summing Up (New York: Random House,
ts78).
4. Eugene Cernan, cited in Frank White, The Oueruiew Efect: Spaee Ex-
ploration and Human Evolution (Boston: Houghton Miffiin Co., 1986), pp.
zo6-2o7.
5. Jacques Lacan, "The Mirror Stage as Formative of the Function of
the I," in Ecrits, trans. Alan Sheridan (NewYork: W. H. Norton & Com-
pany, 1977), pp. r-7.
6. Ilya Prigogine and Isabelle Stengers, Order Out of Chaos (NewYork:
Bantam, r984).
7. R. Swenson and M. T. Tirrvey, "Thermodynamic Reasons for Per-
ception-Action Cycles," in Eeological Psychology 3, no. 4 Q99t): 3t7-
348. See also Rod Swenson, Sltontaneous Order, Euolution, and Natural l-aw
(http : / /members.tripod. com/spacetimenow/contents.htrnl).
8. Humberto R. Maturana and Francisco J. Varela, Autopoiesis and Cog-
nition: The Realization of the Living, Boston Studies in the Philosophy of
Science, vol. 4z @oston: D. Reidel Publishing, r98r).
9. Aristotle, The History of Animals, viii: r, cited in Will Drranq The
Life of Creece (NewYork: Simon and Schuster, r939), p. 530.
ro. 'Willem de Kooning, cited in Richard Marshall and Robert Map-
plethorpe, FtJty New York Artists (San Francisco: Chronicle Books, r986).
r r. James E. Lovelock, "Life Span of the Biosph ere," Nature 296 (r 982) :
56r-563.

247
 248 Notes

CHAPTER 2. LOST SOULS

The epigraphs are from William Shakespeare, Measure for Measure, Act III,
scene one; and Algernon Swinburne, Atalanta in Calydon, in The Poems o;f Al-
(NewYork: Harper & Brothers, r 9o4), p. z 8 3 .
gernon Charles Swinbu rne, vol. 4
r. For the view that humankind is marked by its self-deceptive fune-
real rites and graveyard mysticism, see Robert W. Sussman and Thad
Bartlett, "Deception in Primates," Abstracts of the AAAS Annual Meeting:
199r, AAAS Publication 9r-o2s, Washington, D.C.
z. Fernel, cited in Frangois Jacob, The Logn of Lfe: A History of Hcred-
ity, trans. Betty E. Spillman (New York: Pantheon, r973), p. 25.
3. Descartes to Marin Mersenne, 13 July 164o, in Leonora Cohen
Rosenfield, From Beast-Machinc to Man-Marhine: Animal SouI in French kt
tersfrom Descartes to l-aMettrie (New York: Oxford Universiry Press, r94r),
quoted in Hans Jonas, The Phenomenon of LtJe: Toward a Philosophical Biol-
ogy (NewYork: Harper and Row, r966), pp. 55-j6.
4. Descartes, to Marin Mersenne, cited in Morris Berman, Coruing to
Our Senses: Body and Spirit in the Hidden History of the West (NewYork: Si-
mon and Schuster, 1989), p. 239.
5. Galileo, 11 Saggiatore, tn Opere (Florence, r89o-r9o9) 6: z3z.
6. 'Watts quotation is cited in Michael Dowd, The Big Pkture; or the l-arger
Contextfor All Human Attiuities (Woodsfield, Ohio: Livrng Earth Institute,
r993).
7. Goethe, cited in Thomas H. Huxley, "The Threefold Uniry of Life,"
in S. Zuckerman, Classks inBiology (Port Washington, N.Y.: Kennikat Press,
ry7r), pp. r2-r3.
B. Haeckel, cited in Alfred Russel Wallace, The World o;f Lfe: A Mani-
festation of Creatiue Power, Directiue Mind and Utimate Purpose (London: Chap-
man and Hall, 19r4), p. 5.
9. Ibid., p. 6.
ro. Charles Lyel,, The Principles of Ceology: An Attempt to Explain the
Former Changes of the Earth's Surface, rst ed., vol. z (London: John Mur-
ray, r83z), p. r8i.
rr. Isaac Asimov, Asimov's Biographical Encyclopedia of Science and Tbch-
nology, rev. ed. (Garden Ciry N.Y.: Doubleday, r98z), p. 267.
rz. Alexander von Humboldt, Cosmos: A Skexh oJ a Physkal Descriptiott
of the (Jniuerse, vol. r, trans. E. C. Otte (NewYork: Harper & Brothers,
r85r), pp. 344-345.
r 3 . Theodosius Dobzhansky, "Nothing in Biology Makes Sense Except
in the Light of Evolution," American Biology Tbacher 35 GgZi, rz5-t29.
 Notes 24g

r4. The phrase is from A. V. Lapo, Tiaces of Bygone Biospheres (Oracle,

Ariz.: Synergetic Press, r987).
r5. Vladimir I. Vernadsky, The Biosphere, ed. Mark McMenamin and
trans. David Langmuir (New York: Springer-Verlag, Copernicus, 1997),
pp. 44,58. See also Vernadsky's "The Biosphere and the Noosphere,"
American Scientist 33 094): r-t2.
16. James E. Lovelock's three books on Gaia are Gaia: A New laok at
Lfe on Earth (Oxford: Oxford lJniversity Press, t979), The Ages of Gaia: A
Biography oJ Our Liuing Earth (r ev. ed., New York: Norton, t 99 5), and He al-
ing C aia : Pructical Medicine for the P/aref (New York: Harmony Books, r 99 r ).
17. Wolfgang E. Krumbein, ed., Biogeochemistry oJ Earth, Phoebus and
Titan (Oxford: Blackwell, r983), p. 93.

CHAPTER 3. ONCE UPON A PLANET

The epigraphs are from Cyril Ponnamperuma, The Origins o/ Llft (New
York: Dutton, rgTz), p. r6; Franqois Jacob, The l-ogi of Ltfe: A History of
Heredity, trans. Betry E. Spillman (New York: Pantheon Books, ry73), p.
3o5; and Stanley Miller and Leslie Orgel, The Origins oJ Lfe on Earth (En-
glewood Cli11s, NJ.: Prentice-Hall , ry74.
r. Francis Crrck, Life ltself: l* Origin and Natare (NewYork: Simon and
Schuster, r98r).
z. Aristotle, Parts of Animals (Cambridge, Mass.: Harvard Universiry
Press, r968), Book r, chapter 5.
3. Descartes, cited in FrangoisJacob, The bgn of Life: A History of Hered-
ify, trans. Bery E. Spillman (New York: Pantheon, rg7i, p.
53.
4. Francesco Redi, Esperienze intorno all generazione degli inettis (t668),
cited in Charles Singer, A History oJ Biology (London: Abelard-Schuman,
196z), p. 44o.
5. Redi, cited in Gordon Rattraytylor, The Science of Lrfe: A Picture
History oJ Biology (NewYork: McGraw-Hill, r963), p. I r3.
6. Charles Darwin, Lfe andktters, vol. 3 (London:John Murray, r888),
p.18.
7. A.l. Oparin, The Origin of LiJe, trans. Sergius Morgulis (NewYork:
Macmillan, r938), pp. 247-250.
8. J.B.S. Haldane, "The Origin of Life," frorn The Rationalist Annual
(rSzil.Reprinted in David'W. Deamer and GaiI R. Fleischaker, eds., Orl-
gins oJ Life : The Central Concepts (Boston: Jones & Bartlett, r 994), pp. 7 3-8 r.

9. Cyril Ponnamperuma, The Origins o/Lp (New York: Dutton, r 972),

p.2r.
 2so I
Notes

ro. A. G. Cairns-Smith, Cenetic Thkeouer and the Mineral Origins of Lfe

(New York: Cambridge Universiry Press, r98z).
I L Freeman Dyson, O r igins of Life (Cambridge : Cambridge Universiry
Press, r985).
rz. ErichJantsch, The Sef-OrganizingUniverse: Scientifc and Human Im-
plications of the Emerging Paradigm oJ Euolution (NewYork: Pergamon, r98o),
p.3r.
13. A.D. Ellington and J.'W. Szostak, "Selection in uitro of Single-
Stranded DNA Molecules that Fold into Specific Ligand-Binding Struc-
ttres," Nature 335 $992):85o-852.
r4. lM. Gilbert, "The DNAWorld," Nature 319 (r986): 6r8.
r 5. Harold J. Morowitz, Beginnings oJ Cellular Life: Metabolism Recapitu-
lates Biogenesis (New Haven: Yale (Jniversity Press, r99z), p. 8.

CHAPTER 4. MASTERS OF THE BIOSPHERE

The epigraphs are from Sorin Sonea and Maurice Panisset, A New Bacte-
riology (Boston: Jones and Bartlett, 1983), p. r; and Gordon RattrayThy-
lor, The Science of Lfe: A Picture History oJ Biology (New York: McGraw-
Hill, r963), p. zzt.
r. For discussion of the microfossils of Barghoorn and of Walsh and
other revealers of pre-Phanerozoic stromatolite and rnicrobial wisdom, see
chapters 5 and 6 inlynn Margulis, Symbiosis in Cell Euolution, znd ed. (New
York: Freeman, r993).
2. Bruno, cited in WolfgangE. Krumbein and Betsey Dyer, "This Planet
Is Alive: Weathering and Biology, a Multi-Faceted Problem," in The Chem-
istry of Weathering, ed.J.I. Drever (Dordrecht and Boston: D. Reidel, r985),
P.145.

CHAPTER 5. PERMANENT MERGERS

The epigraphs are from C. Dobell, Antony uan l*euwenhoek and His "Lit-
tle Animali' (NewYork: & Russel, r958); Charles Darwin, The Vari-
Russel
ation of Animals and Plants Under Domestication, vol. z (New York: Organe
Judd, 1868); StephenJay Gould, foreword to first edition of Fiue King-
doms: An lllustrated Cwide to the Phyla of LiJe on Earth, znd ed., by Lynn
Margulis and Karlene V. Schwartz (New York: Freeman, 1988); and Lynn
Margulis, Heather I. McKhann, andLorraine Olendzenski, Illustrated Clos-
sary oJ Protoctlsla (Boston: Jones & Bartlett, r993), pp. ix-x.
 Notes I zst

John Hogg, "On the Distinctions of a Plant and an Animal, and on

r.
a Fourth Kingdom of Nature," Edinburgh New Philosophical Journal tz
(r86r): zr6-225.
z. Ernst Haeckel, History oJ Creation, vol. z (NewYork: D. Appleton,
r889), p.45.
3. Ernst Haeckel, Euolution of Man, vol. I (NewYork: D. Appleton,
r887), p. r8o.
4. David C. Smith, "From Extracellular to Intracellular: The Establish-
ment of a Symbiosis," Proceedings oJ the Royal Society, l-ondon zo4 $g7g):
r r5-r3o.
i. I. E. Wallin, Symbionticism and the Chigin of Species (Baltimore: Williams
&'Wilkins, r9z7), p. 8.

CHAPTER 6. THE AMAZING ANIMALS

The epigraphs are from Charles Darwin, "Difficulties on Theory," in Or'l

the Otigin oJ Species (reprint, New York: Penguin Books, t985), p. zo5;
'William Shakespeare, The Tbmpest, Act I, scene &vo; and Ralph Waldo
Emerson's poem May-Day.
r.
Donald R. Griftin, Animal Minds (Chicago: University of Chicago
Press, r99z), p.253.
z. A. G. Fischer, "Fossils, Early Life, and Atmospheric History," Pro-
ceedings National Academy of Sciences 53 (1965): t2a1-tzr5.

3. Harry B. Whittingtory The Burgess Sfuale (New Haven, Conn.:Yale

Universiry Press, r985), p. r30.
4. Mark McMenamin and Dianna Schulte McMenamin, Hypersea: Life
on ltnd (New York: Columbia University Press, r994).

CHAPTER 7. FLESH OF THE EARTH

The epigraphs are from R. Gordon Wass or,, Persephone's Quest: Entheogens
and the Origins oJ Religion (New Haven, Conn.: Yale (Jniversiry Press,
r986), p. 75 ("entheogen" is'Wasson's term for hallucinogens); and Fran-
ciscus Marius Grapaldus, De Partibus Aedium, book II, chapter 3 (n.p.,
| 492).
r. Jun Takami, cited in Andrey V. Lapo, Tiaces of Bygone Biospheres (Or-
acle, Ariz.: Synergetic Press, r987), p.t2g.
z. Veillard, cited ibid., p. r78.
S.
3. Clive Brasier, 'A ChampionThallus," Nature 356 (rg9z): 382.
 252 Notes

4. Maurice Blanchot, "Literarure and the Right to Death," in Thc Caze

of Orpheus and Other Literary Essays, ed. P Adam Sydney, trans. Lydia Davis
(Barrytown, N.Y.: Station Hill Press), p. 46.

CHAPTER 8. THE TRANSMUTATION OF SUNLICHT

The epigraphs are from William Blake, Sorgs o-f Innocence and oJ Experience:
Shewing the Tbo Contrary States o;f the Human Soa/ (Slough: (Jniversiry Tu-
torial Press of London, rgjS), pp. 2r-22; Georges Bataille, The Accursed
Share, trans. Robert Hurley (New York: Zone Books, r988), pp. r2-r3
(italics are in the original); and Vladimir Vernadsky, The Biosphere (I\ew'
York: Springer-Verlag, Copernicus, 1997), p. 48.
r. Frits W. Went, The Plants (New York: Time, Inc., 1963), p. 16.
2. For a comprehensive view of life in the Proterozoic eon, see Early
Life on Earth, ed. Stefan Bengston (NewYork: Columbia Universiry Press,
r 99-5).

CHAPTER 9. SENTIENT SYMPHONY

The epigraphs are from Erasmus Darwin, Zoonomia; or, the l-arus oJ Organic
Lrf, OlS4; Blaise Pascal, Pensies, no. 93; andLeonardo Tar6n, Parmenides:
A Tbxt with Tian.slation, Commentary, and Critiml Essays (Princeton, NJ.:
Princeton University Press, r965), pp. xx-xxi (the Parmenides quore is a
famous fragment of pre-Socratic philosophy).
r. Clair Folsome, in "Microbes," The Biosphere Catalogue, ed. Tango
Parish Snyder (Oracle, Ariz.: Synergetic Press, r98 j), pp. 5r-j6.
z. Stephen J. Culver, "Foraminifera," in Fossi/ Prokaryotes and Protists,
ed. Jere H. Lipps (Boston: Blackwell Scientific Publications, r9y), p. zz4.
3. 'A Response-Regulated Model in a Simple Sen-
D. E. Koshland, Jr.,
sory System," Science ry6 (tggz): ro55-ro63.
4. The most comprehensive source for these little-known events comes
from Henry FestingJones, "The Butler-Darwin Quarrel," tn Sanruel But-
ler, Author oJ Erewhon (t B j 5-t 9oz): A Mentoir, ed. Henry Festing Jones,
vol. z (London: Macmillan, r9r9), pp. 446-467. The "mental anachro-
nism" quote can be found on p. 447; the "it never occurred to me" on p.
448; and "how the accident arose" on p. 4S3.Butler's fascinatine but
difficult-to-find "evolution books"-tf and Habit; EuolutiLtn, Old and New;
and Luck or Cunning-comprise volumes 4, 5, and 8, respectively, of The
Shrewsbury Edition oJ the Works of Samuel Butler, ed. Henry Festing Jones
and A. T. Bartholomew (NewYork: E. P Dutron and Co., r9z4).
 Notes I zsa

5. Niels Bohr, Physical Science and the Problem o/Lfe (NewYork: Wiley,
r958), p. roo.
6. Samuel Butler, "The Deadlock in Darwinism," in The Humour oJ
forLi-
Homer anil Other Essays, ed. R. A. Streatfeild (Freeport, N.Y.: Books
braries Press, r967), pp.253-254.
7. William Calvin, The Cerebral Symphony (NewYork: Bantam, 1989).
See also GeraldEdelman, Neural Darutinism (NewYork: Basic Books, rg87).
8. For more on the emergence of a superhumaniry see Gregory Stock,
Metanran: Humans, Machines, and the Birth oJ a Clobal Superorganism (Lon-
don: Bantam, r993). For the relationship berween increasingly lifelike ma-
chines and increasingly engineered life, between, as the author says, the
"born and the made," see Kevin Kelly, Out of Control: The Rise oJ Neo-
Biological Ciuilization (Reading, Mass. : Addison-'Wesley, r 994).
9. Samuel Butler, "Darwin among the Machines" and "Lucubratio
Ebria," in The Note-Books oJ Samuel Butler, Author of " Erewhon," ed. Henry
FestingJones (r863; reprint, NewYork: Dutton, t9r7), pp. 46-53.

EPILOG UE

The epigraph is from a letter to Mrs. Holland, c. r88r , quoted in Richard

B. Sewall, The Life of Emily Dkkizsor (NewYork: Farrar Straus and Giroux,
r98o), p. 624.
 Clossary
acanthomorph a rype of acritarch
with spherical vesicle and simple
or branching spires; derived from
Greek acanthe, spiny.
acritarch an organic-walled
microfossil of uncertain origin,
most likely fossil remains of
protists, often spherical.
algae a large group of all the
many rypes of photosynthetic pro-
toctists ranging in size from minute
single cells (<rp- in diameter)
to large seaweeds such as kelp
(> ropm long). The microscopic
floating algae, also called photo-
plankton, form the basis of the
marine food chain. Term includes
chlorophytes (green algae), eugle-
nids, diatoms, and many others,
but excludes "blue-green algae,"
now known to be the rype of
photosynthetic bacteria that
produce oxygen.
amebas single-celled protoctists
that constantly change their shapes.
anabolic characterized by anabo-
lism, the synthesis of complex
molecules from simpler ones. For
example, starch, glycogen, fats, and
proteins are products of anabolic
pathways from sugars, faffy acids,
and amino acids.
anastomosis netforming.
angiosperm a flowering plant; an
organism that belongs to phylum
Anthophyta. It flowers and pro-
duces seeds in a Iluit, a structure
that develops from the fecund
ovary.
Animalia See animals.
animals members of one of the
five kingdoms into which all living
organisms are placed (Kingdom
Animalia). Animal cells are hetero-
trophic and display internal motil-
iry. Animals develop by fertilization
of egg by sperm; gametes are pro-
duced by meiosis. The fertile egg
forms an embryo, the blastula.
apoptosis programmed (geneti-
cally determined) cell death.
asci spore cases of fungi, sexual
structures.
ascofungi fungi that form asci.
ascomycote a phylum of fungi,
molds, morels, yeasts, and others
that are characterized by form-
ing asci, sacs, or capsules when
hyphae sexually fuse and spores
develop as the zygotes undergo
meiosis.
:rscospores spores formed by
ascofungi.
ATP adenosine triphosphate, a
phosphorus-, carbon-, nitrogen-,
255
 256 Clossary
oxTgen-, and hydrogen-containing
ring compound that is universally
used by life to store energy in its
phosphate bonds.
autopoiesis life's continuous
production of itself. Autopoieric
behavior, characteristic of all living
matter, refers to the chemical ac-
tivities (metabolism) of identifiable
organic beings as they self-maintain.
-Without
it they are not alive.
autotrophs organisms that
produce their own food from
carbon dioxide and derive energy
from light or inorganic chemicals,
either by chemosynthesis or
photosynthesis.
axoneme axis or shaft extending
the length of an undulipodiurn,
made of microtubules and other
proteinaceous celi materiai.
bacteria members of one of
the five kingdoms into which all
living organisms are placed
(Kingdom Bacteria, also called
Monera or Prokaryotae). Bacteria
cells are always prokaryotic; they
lack a membrane-bounded
nucleus.
bacteriophage virus in or on
bacteria.
basidia club-shaped sexual struc-
tures of mushrooms and other
fungi.
basidiomycote a phylum of fungi
in which spore-releasing basidia
form: mushrooms, yeast, pufibal1s,
mold.
basidiospores spores formed on
basid'ia.
benthic bottom-dweliing. De-
scriptive of organisms living at the
bottom in aquatic environments
(sea, lake, river).
BIF banded iron formation. A
large sequence of sedinrentary
rocks that consists of iron oxides
and chert occurring in prominenr
layers or bands of brown or red
and black; iron ore.
biophilia love of life.
biosphere the place at the Earth's
sr.rrface. from the upper linrits
in the atmosphere to the lowest
depths in the ocean, where life
exists.
blastula early stage in the devel-
opment of the animal embryo
foilowing cleavage of the zygote:
in many, a fluid-filled, hollow ball
of cells, one layer thick. It is a
unique, defining element of the
animal kingdom.
bryophyte a phylum of nonvascu-
lar, mainiy terrestrial, plants that
includes mosses, hornworts, and
liverworts.
cell material organic compounds
rh:rt make up bodies of living
organisms.
cellulose a sugar-rich compound
of cell walls of plants and some
protoctists.
centriole an intracellular organelle
r X o.25pm in size that consists of
nine groups of peripheral micro-
tubules around a central caviry. A
kinetosome that lacks its axoneme,
this nricrotubular structure forms
at each pole of the nritotic spindle

during division in most animal
cells.
chaetophorales taxon of green
algae.
chemosynthesis the process by
which chemical energy of cells is
generated by inorganic chemical
reactions such as the oxidation of
hydrogen (H:), of sulfur (H2S),
of methane (CH+) and of ammo-
nia (NH:). The energy is used to
reduce carbon dioxide to cell
material.
chitin a sugar-rich, nitrogen-
containing compound of cell walls
of fungi and insect exoskeletons.
chlorophytes green algae.
chloroplasts intracellular struc-
tures, or organelles, in which pho-
tosynthesis occurs. They contain
chlorophylls that are active in
synthesizing starch, protein, and
orher materials (photosynthate).
chromatograph machinethat
separates chemical compounds
that are often colored.
ciliates a phylum (large related
group) of protoctists. Most of
the ro,ooo plus species are rapid
swimmers, single cells with rwo
kinds of nuclei and distinctive
arrangements of their undulipodia.
coccolithophorids protoctists
belonging to the phylum Prymne-
siophyta that produce decorative
calcium carbonate scales (cocco-
liths). Most are marine algae; many
are known from the fossil record.
conidia fungi molds that produce
spores without a sex act.
Clossary 257
conidiospore spores formed by
conidia fungi.
coPepods a class of crustacean
animals lacking a carapace and com-
pound eyes. Their first thoracic
appendages are modified for "filter
feeding," removing food particles
from the surrounding water.
cross walls cell walls.
cryptozoans a term first used
by geologist Charles Walcott
for what he thought were some
strange animal; now known to
be Cambrian stromatolites.
cyanobacteria oxygenicphoto-
synthetic (photoautotrophic)
green-pigmented bacteria.
Sometimes still called plants
or "blue-green algae," they are
neither. They evoived when
photosynthetic bacteria, employing
the unique chlorophyll system,
mutated from purple predecessors
to get their hydrogen atoms from
water. More than ten thousand
kinds have been catalogued. It is
believed that they released the
o\ygen that transformed Earth's
atmosphere.
rycadofilicales "seed ferns," pteri-
dosperms: now extinct, abundant
inhabitants of the forests of the late
Paleozoic and Mesozoic eras, 345
to 65 million years ago; the cell
material of their bodies provided
much of the coal deposits mined
today.
cytoplasm the fluid of cells out-
side their nuclei.
deuteromycote the phylum of
fungi that lack sexual stages and
 258 Clossary
therefore cannot be assigned to
either the ascomycota or basid-
iomycota. They reproduce by
release of conidia (a kind of prop-
agule). These thin-walled cells
often form and then break offfrom
the ends of ordinary hyphae.
diploid pertains to a cell or
organism in which each chromo-
some is present in rwo copies. In
animals, all cells are diploid except
for gametes, which are haploid. In
fungi, all cells are haploid except
those that produce spores.
dissipative system nonbiological
patterns formed by energy flow.
DNA deoxyribonucleic acid:
a long chain molecule made of
organic compound links called
nucleotides (bases, deoxyribose
sugar, and phosphoric acid) in the
nucleoid of prokaryotes or the
nucleus of eukaryotes. DNA stores
in its nucleotide sequence the
genetic information for making
proteins in all cells.
endolithic Iiving inside rocks.
endorphins with ekephalins, one
of ru/o naturally produced opium-
like compounds in the brain.
endosymbiosis a relationship in
which a member o[ one species
lives notjust near or even per-
manently on a member of another
species, but inside it. Together they
(the bionts) form the symbiont in
a holarchy.
enkephalins See endorphins.
entropy a measure of the amount
of energy in a system that is un-
available to do work, generally in
the form of useless heat, noise, or
uncertainty; the randomness in the
structure of a natural system.
eukaryotes organismsmade
of nucleated cells-cells with a
membrane-bounded nucleus that
undergo some form of nritosis.
Eukaryotes all derive from and
include protoctists; bacteria (pro-
karyotes) do nor. Protoctists,
animals, fungi, and plants are
all eukaryotes.
foraminifera commonly called
forams; marine organisms having
pore-studded shells, or tests,
microscopic to several centi-
meters in diameter. The major
class in the Protoctist phylum
Granuloreticulosa.
forb broad-leavedplant.
fractal in mathematics, any
of a class of complex geometric
shapes that commonly exhibit
the properry of self-sinrilariry. A
self-similar object is one whose
component parts, when sufiiciently
magnified, resemble the whole.
Fungi one of the five kingdoms
into which all living organisms are
conveniently and unambiguously
classified [or acadenric discussion;
also called the Kingdom Mychota.
Fungi are osmotrophic heterotrophs
composed of eukaryote cells and
propagated by spores.
Gaia over thirry million types
of live beings, descendants from
coffinon ancestors, and members
of five kingdoms that produce and
remove gases. ions. and organic

compounds. Their interacting
activities lead to modulation of
Earth's temperature, acidiry and
atmospheric composition.
gall insect, virus, or fungal-
induced usually spherical structure
on plants.
gametangia sex organ that
contains gametes.
gametes haploid cells of some
protoctists, nearly all animals and
many plants, capable of fusion; i.e.,
cells that fuse (fertilize) to produce
a diploid zygote to harbor double
(zN) the number of chromosomes.
In animals, the zygote (fertilized
egg) develops into a new diploid
organism. In plants, the gametes
are produced by adult haploid
organisms (called gametophyte$
by the process of mitosis.
gametophyte plant with cells that
contain one set of chromosomes.
gastrula embryo stage of animals
as their digestive tract is forming.
genome the total genetic make-
up of an organism. This includes
dl the native DNA in each cell.
gymnosperm plant in
a seed
which the ovules are carried naked
on cone scales. The most familiar
groups are the conifers (pines,
spruces, firs, etc.).
haploid pertains to a cell or
organism containing in each cell
only one representative of each
chromosome. (See also diploid.)
hemoglobin oxygen-carrying
blood protein.
Clossary 25g
heterocysts enlarged and special-
ized cells in cyanobacterial fila-
ments in which nitrogen fixation
occurs. This process makes avail-
able to living matter the inert
nitrogen (N2) so plentifirl as a gas
in the atmosphere and so essential
to protein formation.
heterotrophs organisms unable to
produce their own food and unable
to use light or inorganic chemical
energy; they obtain their energy
and carbon, nitrogen, and other
essential elements from organic
compounds produced by primary
producers (chemosynthesizers or
photosynthesizers).
holarchy word coined by Arthur
Koestler to express the coexistence
of smaller beings in larger wholes
and avoid the value-ridden term
"hierarchy." The constituents he
named holons-wholes that also
function as parts.
holon a part or component of
a holarchy; see above.
humus organic-rich soil.
hypha pl. hyphae. The filaments
or thin tubes that comprise the
web structure of fungi.
integument any protective body
covering, such as cuticle or skin;
in plants, the covering of the ovule
that develops into the seed coat.
iscedia lichenpropagule.
kinetosome anintracellularorgan-
elle, not membrane-bounded, char-
acteristic of all undulipodiated cells.
Microtubular strucrures necessary
for the formation of all unduli-
 250 Clossary
podia, kinetosomes differ from cen-
trioles in that from them extends
the shaft, or axoneme.
Kingdom highest taxon, which
contains smaller groups (phylum,
class, order, family, genus, species)
of closely related organisms
classified on the basis of body
form, genetic simiiariry metabo-
lism (body chemistry), develop-
mental pattern, behavior, and
other characteristics. See a/so
animals; bacteria; Fungi; plants;
Protoctista.
laboulbenomycetes a kind of
ascofungus.
lichens association of fungi with
algae or cyanobacteria, which
provide photosynthesis. There
are an estimated z5,ooo different
kinds.
lignin the chemically complex
polyphenolic ciass of organic
compounds that give plants their
woody hardness.
lithosphere rocky surface of
Earth, including the regolith
continental rocks and ocean floor.
magnetotaxis attractionto
magnetic pole or force.
mats an ecological term for an
aquatic communiry of bottom-
dwelling organisms such as
cyanobacterial-dominated microbial
mats, a precursor to stromatolites.
meiosis one or rwo successive
divisions of a diploid nucleus in
which the number of chromo-
somes is reduced by half, leading to
haploid offipring (containing half
the genetic complement of the
parent cell).
metabolism the sum of the chem-
ical and physical processes that
occur in all living organisms and
involve incessant replacement of
their chemical constituents.
microbes (microorganisms) beings
best seen through a microscope.
micrometer (pm) one millionth
of a meter.
mitochondria intracellular cell
organelles, the sites of ATP syn-
thesis, probably began as oxygen-
respiring purple bacteria. They
were incorporated with others
through symbiosis to create new
kinds of cells. Today mitochondria
persist even in human cells as
organelles that use oxygen to per-
form respiration and generation
of chemical energy. They are thus
examples of holons. See hoiarchy.
mitosis the division of the
nucleus, characteristic of eukary-
otic cells, that produces two
offspring cells; ploidy of the
nuc'leus is unaltered by nritosis.
Monera another name, along
with Prokaryotae, of Kingdom
Bacteria. Comprises alJ organisms
with prokaryotic cell structure:
they have small ribosomes sur-
rounding their nucleoids but lack
membrane-bounded nuclei.
mycelium the mass of hyphae
(usually underground) that are
associated as a single fungus
organism.
mycoplasm a group of wall-less
pleiomorphic bacteria.

mycorrhiza symbioticfungi
associated with plant roots. They
assist in the mineral nutrition of
the plant.
natural selection the fact that
more organisms are produced than
can ever survive. Those that survive
to have offspring are said to be
naturally selected.
necrobes organisms that live on
the dead or on dying life.
negentroPy thermodynamic
concept: the recovery of energy
in the form of organization rypical
of life, at the expense of solar or
chemical oxidation energy. (Cf
entropy.)
nematodes roundworms.
neuropeptide a natural compound
(like protein) that commu-
a small
nicates behveen nerve cells.
noosphere the global nrind; the
aggregate net of throbbing life and
its consciousness surrounding the
Earth.
nucleated characteristic ofplant,
aninral, fungal, and protoctistan
cells, those that bear nuclei as their
defining feature.
nucleic acids long chain mole-
cules like DNA or RNA.
organelles distinctstructures
within eukaryotic cells that are
distinguishable by microscopy.
Exanrples incltrde plastids, mito-
chondria, and nuclei.
ortanic refers to chemical
composition made of carbon and
hydrogen compounds often, but
clossary I zel
not necessarily, made by living
organisms.
organism any living being,
member of one of the five
kingdoms of Life (thus viruses
are not included).
osmotrophy a mode of hetero-
trophic nutrition where small
organic chemical molecules are
absorbed through cell walls, if
present and actively transported
across cell membranes. Fungi, most
bacteria, and many protoctists are
osmotrophic.
parthenogenesis development
of an egg without fertilization
by sperm.
pathogen any organism that is ca-
pable of causing disease or a toxic
response in another organism.
phaeoplasts brownplastids,
photosynthetic organelles of
phaeophytes.
photosynthesis the process by
which chemical energy is gener-
ated from sunlight: certain hexose
carbohydrates are formed from
carbon dioxide and water in the
chloroplasts of living plant cells,
with oxygen or suifur produced as
a waste product from compounds
that provide the hydrogen (i.e.,
water, hydrogen sulfide).
phylogeny family tree diagram.
planktic (planktonic) descriptive
of organisms in the water column
moved by currents.
Plantae See plants.
plants members of one of the
five kingdoms into which all living
 262 Clossary
organisms are placed (Kingdom
Plantae). Plants are multicellular
eukaryotes, generally rooted in the
earth or on other plants (epiphytes)
Most make their own food by pho-
tosynthesis. although rhis is not a
defining characteristic. Plants all
grow from spores at one stage of
their life cycle and from maternally
retained embryos at another.
plasmid a small replicon, or short
piece of DNA, that occurs natur-
ally in bacteria. It can enter into
the ceils within roots and stems
of susceptible piants, bringing bac-
terial genes into the plant's nuclei.
plasmodesmata connections
between plant cells, cytopiasmic
strands that extend through
perforations in the cell walls.
plastid the generic term for
photosynthetic organelles in plants
and protoctists (all algae). Bounded
by double membranes, plastids
contain the enzymes and pigments
for photosynthesis, ribosomes,
nucleoids, and other structures.
ploidy the number of chromo-
some sets in the nucleus of eu-
karyotic cells (e.g., diploid refers
to rlvo sets; triploid to three).
polychaetes a class of the phylum
Annelida. These animals, the
paddie-footed worms, nearly all
dwell in marine habitats.
progymnosperms a large group
of early, extinct plants ancestral to
modern fornrs.
prokaryotes bacterium; member
of the Kingdom Bacteria (King-
dom Monera or Prokaryotae); cell
or organism composed of cells
lacking a membrane-bounded
nucleus (nucleoids).
propagule any cell or multi-
cellular structure produced by
an organism that is capable of
survival, dissemination, and
growth. Propagules include all
the sexual and asexual methods
of reproduction, survival, and
propagation.
prophage a piece of a bacterial
genome, a small replicon (DNA)
that has temporarily integrated into
the rest of the bacterial genome.
proprioception the detection of
self.
Protists nricroscopic protoctists,
either single-celled or composed
of very Gw cells.
Protoctista one of the five king-
doms into which all living organ-
isms are grouped for academic dis-
cussion. Protoctists are eukaryotic,
nucleated microorganisms (the
single-celied protists) and their
immediate multicellular descen-
dants. The kingdom includes all
eukaryotic organisms with the
exception of animals, plants, and
fungi; for example, all algae, slime
n-rolds, amebas, slime nets, water
molds, and foraminifera-an
estimated 2Jo,ooo extanr species
in about fifry major groups.
psilophytes an extinct group of
early plants that lack leaves and
flowers.
regolith loose covering of rocky
material at the surface of any
planet.

replication the process that occurs
when a structure (e.g., a DNA
molecule, a crystal) produces a
second structure exactly like itself.
replicon piece of DNA capable
of replication.
reproduction process by which
living cell or organism produces
another very sirnilar being. The
second being may differ because of
mutation, genetic recombination,
symbiotic acquisition, developmen-
tal variation, or other factors.
rhodoplasts red plastids, photo-
synthetic organelles of all rhodo-
phytes (red algae), some cryp-
tomonads and other protoctists.
ribosome cell organelles present
in large numbers and involved in
the synthesis of protein in cells of
all organisms.
RNA ribonucleic acid: a long
chain molecule made of organic
compound links called nucleotides
ftases, ribose sugar, and phosphoric
acid), one kind of which translates
the sequence information in DNA
into the sequence of amino aci& in
proteins in the cytoplasm of all cells.
sclerites patterned fossils of tiny,
shelly plates of unknown origin
present at the beginning of the
Cambrian Period, 54r rnillion years
ago, in many sedimentary rock
deposits worldwide.
sessile descriptive of an animal
or protoctist that lives permanendy
attached to substrates-e.g., sea
sponges or benthic foraminiGra that
are attached to rocks (6[ vagile)-or
a plant structure direcdy attached
Glossary 263
to the main body of the plant
rather than to a secondary stalk.
soredia lichenpropagules,
photobionts interwvined with
fungal mycobiont.
spectroscope a machine that
a analyzes compounds using light.
spicule any hardened, pointed
body part; for instance, the hard
internal structures in sponges
composed primarily of sfica or
calcium salts.
spirochete motile heterotrophic
bacterial cells in which flagella
inserted at the ends of the cell
are wound beneath the inner and
outer membrane in the flexible
cell wall.
spores propagulesusuallyresistant
to extreme conditions.
stromatolites domed and layered
rocks formed by the remains of
communities of bacteria and
minerals that trapped, bound, and
precipitated materials in an ordered
way.
symbiogenesis an evolutionary
term referring to the formation
of new life forms, new organs, or
new cell organelles by permanent
association of older, preestablished
life forms.
symbiosis an ecological and
physical relationship between wvo
or more kinds of organisms such
that they live together. These
merging partners are called
symbionts.
synergy interactionberween
entities that together behave as
more than the sum of their parts.
 264 Clossary
Life, love, and social behavior
appear to be synergistic phenom-
ena, as does the behavior of
organisms formed in permanent
symbiotic combination.
taxon pl. taxa. A group of living
organisms, ranked ftrom species to
kingdom (see under Kingdoms).
tetrapods four-legged animals.
thallus leaflike bodies of cyano-
bacteria, algae, and certain plants
(liverworts, mosses) that do not
make flowers.
thanosis the process, internal to
an organism, of aging and dying;
apoptosis, or programrned cell
death.
thermodynamics the science of
energy states and flows. The first
law of conservation of energy says
that energy (chemical, kinecic, nre-
chanical, heat, sound, wave, etc.)
may change form and become less
available for work, but it does not
diminish in quantiry. The second
law of thermodynamics says that
the qualiry of energy diminishes
over time and that energy's pro-
pensiry to dissipate leads to natur-
ally arising organized energy-
dissipating structures, of which
life is one example.
thermoplasma bacteria that live
in high-temperature, acidic, and
usually sulfur-rich environments.
They lack cell walls and are
extremely variable in shape.
trilobite group of extinct
a great
marine arthropods that thrived in
the Paleozoic Era.
undulipodium p/. undulipodia.
Motility organelle that shows
swimming, feeding, or sensory
function; composed of at least 2oo
proteins and always underlain by
the centriole-kinetosome from
which it develops. Its [9(z)+z]
rnicrotubular shaft, the axoneme,
is covered by a cell mernbrane and
so-called flagella of eukaryotic
cells. Examples include cilia, sperm
tails.
vagile characterized by the abiliry
to move freely (l sessile).
virus an infectious agent that
replicates only inside living cells;
not autopoietic. An organized set
of functioning chernicals, capable
of replication but not of any
metabolism and therefore not a
unit of life. A protein-coated
plasmid is one description.
zygofungus a mold that mates to
form spore cases called zygospores.
zygomycote a phylum of fungi
characterized by joining and fusing
open hyphal tubes through which
nuclei flow andjoin. The zygo-
spore (really a sporangium) results
when different complementary
hyphae fuse and their nuclei fer-
tilize and undergo meiosis.
zygote a fertile egg; the diploid
nucleus or cell produced by the
fusion of rwo haploid nuclei or
cells in fertilization. The first stage
in the embryo of animals and
plants.
Acknowledgments

The revival of Schrcidinger's inquiry into "What is life?" was Peter

N. Nevraumont's idea.'We are grateful to him and to the many oth-
ers who helped realize this work. Michael Dolan collected illustra-
tions and critical conlments, prepared the timeline, checked dates
and references, and made numerous other invaluable contributions
to manuscript preparation; Connie Barlow gave us many useful sug-
gestions as she edited the manuscript, including her lead to Rod
Swenson's missing link between thermodynamics and mind. We
thank Curt Staeger of Paul Smith's College of Arts and Sciences in
New York for sending us the tape of his radio show on the subject,
and his student Deborah Smith for her statistical survey ("I wanted
to research this topic because I want to know what life is. The way
I gathered my information was by writing to scientists. . . . From
the information I collected, it seemed like no one really knew what
lift is. Rather, they chose to tell me the physical features of life, or
offered analogies and metaphors. Most students answered by sin-
gling out humans specifically"). Thanks to David Abram for his
never-say-die animism and his close reading of the rnanuscript. We
are indebted to Osborn Segerberg, Jr., for sharing his research and
experiences. (The secretary of James 'Watson, codiscoverer of
DNA's structure, replied with unintentionally dry humor to
.Watson
Segerberg: "l am so very sorry, but Dr. is unable to answer
'-What 'With meetings now in
your question, is life?' our surrurler
full swing, his life is most hectic, and he doesn't have time to spare
for anything else.") We are grateful to David Bermudes for his em-
phasis on RNA over exotic cuisine and to Oona Snoeyenbos-'West
for home-ground coffee, foraminifera facts-on-ca11, and instant in-

265
 265 ] nckno*ledgments

dexing. We thank Simone Nevraumont and AnnJ. Perrini for their

expertise at Nevraumont Publishing. Seth! Leary and Bob Hanie and
his students are acknowledged for excitement and hospitaliry Landi
and Rufus Stone for companionship and conversation, Greg Hin-
k1e for the phrase "symbiosis seen from space," Tobi Delbruck for
manuscript review and Caltech skepticism, and Emil Ansarov for
-We
suggesting the need to examine life's shadow death. are espe-
cially grateful to Mary Catherine Bateson, David Bermudes, Free-
man Dyson, Ricardo Guerrero, Donna Haraway, Bryce Kendrick,
Steven Rose, Michael Rothschild, Jan Sapp, James W. Walker, and
'Wilson,
E. O. all of whom made suggestions for inrprovement.
'We
note the contribution of Reg Morrison, whose book The Spirit
in the Gene: Humanity's Proud Illusion and the l-aws of Nahre (Cor-
nell lJniversity Press, r999) extends our thesis but arrived too late
to be incorporated into this text.
We thank the great minds who have dug before us and left us
their shovels and whom we would, in accordance with our post-
structuralist notion of a generalized plagiarism, no doubt credit com-
pletely were it not for our craving for self-aggrandizement, proba-
bly itself an evolutionary legacy. Hans Jonas's brilliant hisrory of
materialism, The Phenomenon of Life, for example, profoundly
influenced us. 'We are very thankful for Eileen Crist's early review
of the Butler manuscripts and for Carl Sagan and Ann Druyan's last-
rninute criticisms and encouragement. We acknowledge the grace
of Christie Lyons's illustrations, some of which were based on pho-
tomicrographs of organisms taken by David G. Chase, William
Ormerod, and Lorraine Olendzenski and published others. The rhi-
noceros beetle came on loan from the department of entomology
at the University of Massachusetts, Amherst. We thank so many
of our colleagues, acknowledged on page 269, who generously pro-
vided illustrations. Fern Serna (Newell Color Lab) and Steve Ger-
ard (Photo Researchers) both went the extra rnile to help us obtain
the best images. We greatly appreciate the aid of Donna Reppard,
Karen Harney, Thomas Teal, Dianne Bilyak, Aaron Haselton, An-
 Acknowledgments I Zel

drew'Wier, and Lorraine Olendzenski in the innumerable tasks re-

quired for manuscript completion.'W'ithout the rapid excellent typ-
-We
ing of Karen Nelson we never could have finished on time. are
grateful to Ralph G. Paul for conceiving and compiling the first draft
of the glossary.
IJnprecedented financial support for the research on which this
project is based came, in part, from the Richard Lounsbery Foun-
dation, NewYork Ciry. We thank Marta Norman for the confidence
she has shown in our scientific communication efforts. The Exobi-
ology and Planetary Biology Internship programs of NASA (Michael
A. Meyer) and both the College of Natural Sciences and Mathe-
matics (deans Frederick L. Byron and Linda Slakey) and the Geo-
sciences Department (Raymond Bradley and Martha Beckwith) at
the Universiry of Massachusetts, Amherst, also came to our finan-
cial aid. We thank Carolyn Reidy, Dominick Anfuso, Patricia Lea-
sure, Linda Cunningham, and'William Rosen (Simon & Schuster)
as well as Anthony Cheetham, Michael Dover, and Lucas Dietrich
(Weidenfeld and Nicolson) for their enthusiastic support of this
book.
We thank Rose Vekony, YukiThkagaki, Howard Boyer, and Char-
lene 'Woodcock for meticulous care of the redesigned paperback edi-
tion. Jos6 Conde is acknowledged for his design of the book's origi-
nal edition. The design of this paperback edition is by Nola Burger.
Any comment we might evoke ftom you, the reader, and espe-
cially you, the science student, should be directed to us at Science-
writers, PO. Box 67 r, Amherst, MA 01004-067 r. Your written re-
sponse to our prose or illustrations is warmly welcomed as the
"what-is-life?" dialogue continues.
 Sources of
lllustrations

PLATES

ra. NASA I3b. Brian Duval, University

of Massachusetts
rb. CNRI/Science Photo Library
r 3 c. Walker/Photo Researchers
z. Johannes H. P. Hackstein,
Universiry of Nijmegen, The r4. Walker/Science Photo Library
Netherlands
r5. Charles Cuttress
3. NASA/Science Photo Library
r6. Vivian Budnik, Universiry
4. David Deamer, Universiry of Massachusetts
of California, Santa Cruz
r7. Electron micrograph by Richard
5a. James A. Shapiro, Universiry Linck, (Jniversiry of Minnesota;
of Chicago false color by David Gray, Woods
Hole Oceanographic Institution
5b. Arthur Winfree, Universiry
of Arizona, Phoenix r Sa-b. J. W. Hastings, Harvard
6. Hans Reichenbach, Braun- Universiry
schweig, Germany r9a. Jukka Vauras
.William
7. Norbert Pfennig, Universiry r9b. Ormerod
of Konstanz, Germany
zo. Biophoto Assoc./Photo
8. Sgepko Golubic, Boston Researchers
Universiry
z r. Connie Barlow
9. Elso S. Barghoorn (deceased)
zz. Jererry Pickett Heaps, lJniver-
roa. Stan-ley Awramik, Universiry siry of Melbourne, Australia
of California, Santa Barbara
23. Jim Frazier, from The Flowering
rob. Peter Westbroek oJ Condwana by Mary White
r r. Tore Lindholm, Abo Akrd..ni,
24. David Mulcahy, Universiry
Finland
of Massachusetts
tz.Lynn Margulis 25. William Ormerod
r ja. Franqois Gohier/Science
26. NASA
Photo Library

269
 27O I Sorr.", of tllustrations
FIG U RES
r. UPl/Bettman Archives
z. Redrawn fromJeremy Sagan,
Cornell Universiry
3. Charles Keeling, Universiry
of Hawaii, and Lloyd Simpson,
Center for the Study of the
Environment, Santa Barbara
4. Mary Beth Saffo, Arizona State
(Jniversity, Tempe
j.
Peter W'estbroek, Universiry
of Leiden, The Netherlands
6. L. CardlScience Photo Library
7. Christie Lyons
8. Lynn Margulis
9-ro. Christie Lyons
I I. Christie Lyons, based on
micrograph of David John,
Oral Roberts Medical School
rz.Prof. FrankE. Round, Uni-
versiry of Bristol, England
r 3. Christie Lyons
r4. R. O. Schuster
r j-r 7. Chrisfie Lyons
r 8. John F. Stolz, Duquesne
University, Pittsburgh
 lndex
Page numbers in italic denote illustrations; color plates, which follow page r44,
are listed by plate number (r.g., pl.r for the plate; pl. [cap.] for the caption).
acquired traits, inheritance of, z3o.
See a/so purposefulness; symbiosis
aerobic metabolism,85, r3o, r33
aesthetics, 30, 3I, 43, pl. z5 (cap.)
African Queen, The, zr7
Agaricus brunescens, 18 5
aging,9r, 97, ro4, r37
agriculture, r35, r 84-r85, zo8-2o9,
2ro, 2r r
Agrobacterium, t8t
alantne, Tz
alchemy, 7
alcohol, 25,74, r88
algae: and atmosphere, r4z; in desert
crusts, 2o2-2o3; pigments and,
r34, pl. tja-c; as plant ancestor,
r7g, rg4, r9J-r96, zo3; plastids
of, rzt; reproduction of, r56,
pl. zz;symbiosis and, roo, rzo,
r27, r33-r34, r36, r77-r78,
pl. t t, pl. t4
Alke in Wonderland, rz7
alpha-androsterol, r 85
Altman, Sidney, 84
Aluin, 174
Amanita, r77, t86
Amazon rain forest, zo8
amebas: reproduction of, r37-r38,
r4r; symbiosis and, r3z, t33
American Revolution, 4o
amino acids, 82, to7
amitochondriates, tz8
ammonia, 70, 7 3-74, 8r, to7
Amoeba proteus, 132, zt8, zz4
t
anaerobic metabolism, 8J, 89, ror-
to3, rz8, pl. 7
Anaxagoras, 59
ancestor worship, 36
Andronicus of Rhodes, 4z
angiosperms, 2o7-2o9, pl. 24, pl. z5
animalcules, 65, 87, r r3
animals: attitudes toward, 37-38. r49-
| 5c, 22o: autopoiesis and, r 5 r,
r55; bacterial dependence of,9r,
g2-g3, g7i behaviors of, r47-r49,
zr4; blastula of, r45, t46, r5z-
r54; deception and, t6S, zzr;
evolution of, r5r-r52, r54-r56,
r59-t64, 167-17o, zo7-2o8, zo9,
pl. t 5; food procurement by, r 5o-
r5r; fungi and, r83-r86; as hetero-
trophs, roo; human devaluation
ol r49-rjo; humans (see humans);
askingdom, rr8, r4r, zr4;mind
and, I5o, r5z, 167-17o, zr4;
mineral production by,25,27, z8-
29, r63-t64, 166-167; minimal
form of, r55;mitosis and, rz3,
126, r28; multicellulariry and,
r35-r36; phyla of, r46, r+7,15r,
r 5 3 ; plant-animal interdependence,
2oo-2o r, 2o7 -2 r o ; plant-animal
split, rr8, tg7, rgg; as preceding
plants and fungi, r47; protocrists
as ancestors of, r33, r4I, r58, I60;
recogrition, of self and others, r48;
reproduction of, 94, r5r, r55, r58;
spontaneous generation of , 6 4-69
271
 272 lndex
animism, 3, j, 8, r7, 33, 4r, 2r7
Annelida, r67
annelid worms, r6o
anthrax,8T-88
antibiotics, g6, roo, r8z, r88
antibodies, r20
Antiquity oJ Man, Thc (Lye11), 46
ants, r68, r84, r8j
apes, r48, 165, z16
apoptosis (thanosis), r37. See also
aging; death
archaebacteria, 89, roo, 1o3, r30, r67
archacprotists, rzz, tz8
Archean eon, 6r, 66-70,84-85,9j,
99, rO2, r04, rro, r3J
archegonium, r96
Aristotle, zo, 35, 36, 42, 64,65
Armillaria bulbosa, 174 175
Armstrong, Neil, z4o
Arrhenius, Svant6, 59-6o
art, 3 r
arthropods, r6r
artificial selection, 22o-22r
ascomycotes, I76
Asimov, Isaac, 46
atheism, J2, 70
atmosphere : biological contributions
to, r2-r4, 20, 2 1, 89, go, 92;
origin of liG and, 7o-74, 89;
oxidation of minerals and,
roj-r06, rz9, r6o; oxygen
(see oxygen); ozonc'layer, ro6,
protoctists and, r4z. See also
biosphere; Earth
NlP,7rt7z, ror, r02, r3o, r8r
Atsatt, Peter, r79
Atta, r84, t85
Augustrne, 64
autopoiesis: animals and, i5r, r55;
biosphere and, zo-24, z t, zq,
5z-54; death as failure of,78-79;
definition of, r7-zo; evolurion
and, lor; global system (Gaia), xiv,
46,52 55, r68-r7o, r89, r90,
pl. ta, pl. z6; nretabolisnr and, 17
18. t9.77 79: rrrind and. zzz;
origins of life and,77 78; plants
and, zrz; purposefulness and,
16-17, 5z-53; vs. reproduction,
r8,43,79-8o; RNA and, 83-85.
See a/.so evolution; reproduction
axonente, pl. t 7
Aztecs, 3 5
bacteria (Monera), 58-60, g7-g8,
t t5, pl. rB; archaebacteria, 89,
roo, r03, r30, 167; attitudes of
Gar or disdain toward, 58, 59, 88,
90-9r. r r 8. .:.r9-zzol autopoiesis
and, r8, r9; discovc.ryof,8Z 88;
disease and, 87-88, roo; DNA
of , 89, 94-96; as endosymbionts,
ideal. rzo: evolution of. 9o, 9.3,
97-gg, r 54; Grmenting, 99-roo,
ror; fossils of, 88-8q, ro7; fur-rc-
tron of, 59, gr-93, p/. 8; gene
trading by, 93-96, 95,97, trr,
r16; as kingdom, rr7, rr8; life
originating as, 79, 90, r ro-r r r,
r59-t6o; locomotion oC gz,
ro2, 122-r23: mat structures
of, 89, ro5, ro9-rro, zoz, pl. 9,
pl. t oa-b; metabolism and, 9r-93;
mind and, zt9-zzo; mineral use
and production by,27, z8-29,
r02, ro9-r ro, 237-238; r'nulticel-
lulariry and, go, pl. 6; and nuclei,
lack of,94; origins of, 58, 59-6o,
89; pollution and recycling by, ro6-
I08; as prokaryotes, 94; reproduc-
tion of, 79-80, 9r, 92, 9+-96, g5;
RNA ot, 89; srze of, 9z; space ex-
ploration and, z4o; sponraneous
gcneration of life theories and, 65,
67-69; stromatolitcs, ro9 r ro,
pl. S, pl. i oa-6; synrbiosrs ancl
(-see symbiosis); as unit of life,
minimal, r8, 58-59, tlz
bactcriophage, 96
Barghoorn, Elso, 88
bariunr, z8
Bar-Nun, Akiva, Zr
basidionrycotes, t76-177, t84,
pl. t 9a-b
 lndex 273

Bassi, Agostrno, 87 brain, r56-157, 23o, 233-234

Bataille, Georges, rg3, rgg-2oo, Brasier, Clive, r75
breath, of biosphere, 23-24, 24,
Bathybius haeclelii, r t8 35-36
bats, r47 Bruno, Giordano,37, S4-SS, gl
Bdellodbrio, t3t bryophytes, zo2,2o4
beaury. See aesthetics bubbles, 75
bees, r4g, r5r, r68 Buddhism, zr8
beedes, 97, t46,pl. z Buffon, Georges Louis Leclerc, 68,
Beggiatoa,59 225
Belousov-Zhaborinsky reaction, 76, Burgess shale, r6o-r63
Pl'sb Butler, George, zz3
Bible, 45-46, 5r,66, 187 Butler, Samuel, 222-229, 23o, 23 t,
big hair, 165 233, 234, 239-240
biophilia theory, pl. 25 kap.) butterflies, r S S, 164-165
biosphere: age of, 5; as alive, xiv, +6,
5z-55, t68-17o, r89, t9o, pl. ra, Cairns-Smith, Graham, 7 4-7 5
pl. z6; artificial, z4z-243; as auto- Calatrava, Santiago, z4r
poietic, 20-24, 2 1, 24, 52-54; es calcium, 26, 27, 28, I09, I ro,
closed system, I9o-I9r, 2ot-2o2, r63-r64
241-2+3; competition vs. coop- Calvin, William,233
eration in, 23, r68, 2ro,2r+,236, Cambrian fossils, 63, 77, t52,
238; energy economy of, r9o- r 59-t64
t9r, r99-2O2, 2ro-2r 2, 2t 5, Campbell, Susan, r95
23 +-23 5, 242, 246:' fungi and, cannibalism, r3g-r4r, | 44
17+, 1 8 1- 1 8 2; reproduction of, capitalism,235
242; temperature maintenance of, carbon: early Earth and, 6r; fungi
22, 23, 32,52; as term, 50; third and, r 8z; interstellar space and,
dirnensional development of, tg4, z5; life origins and,7o,7r,7S,89;
196. See also atmosphere; Earth photosynthesis and, 9o: precipita-
Biosphere z, z4z tion of, 167
Biosphere, TTre (Vernadsky), jo-S r carbon dioxide: atmosphere and, zo,
biotechnology, 93, 96, r8t, 2og 23-24, go, 2o2, 2tti autopoiesis
birds, r47-r48, r58, I67 and, zo, z r , z 4; origins of life
Black Plague, 37 and, 89; photosynthesis and, 9o,
Blake, William, I93, r99 ro2
Blanchot, Maurice, I89 Carboniferous period, zo6
blastula, r 45, t 46, 1 S2-r 54 carbon monoxide, r 3
blood,6,98, r53,228 carnivores, r99
blue-green bacteria. See cyanobacteria Carnot, Nicolas, r5
blue whale, rg4 carotene, t66, pl. tjc
Bogart, Humphrey, zr7 Carroll, Lewis, Iz7
Bohr, Niels, zz7 catalases, r2g
bones, 25, 27, 163-164, 237 Cathedral of Saint John the Divine,
Bonnet, Charles, 66 24r
bowerbirds, t 47-r 48, r 5r Cech, Thomas, 84
brachiopods, r6o Cellarius, 239-24c.
 274 lndex

cell(s): deathof, r J6-r j8; differentia- ClassiJrcation of l-ower Organisms, The

tion of tissue and, r35, r4o, r j3-
r54, rj5; offungi, r72;rnetn-
branes of, 8J-86, roo; mind of,
2rg-22o, zz8-zz9; nuclei of, 94,
rr4, 115,116,r29,i35;ofplants,
r99; as unit of life, minimal, r 8,
78, 82, 85-86; walls of, roo, r72,
2O3
Cenozoic er:, 62, 80,98, 167
centrioles, rz3, rz6-t27, tzg, r57
Cernan, Eugene A., ro
chaetophoraies, zo3
Chagas'disease, r4r
Chatton, Edouard, rr4
cheese, 186, r88
chemotactic bacteria, zt9
chimpanzees, 148,216
China, 35, r87
Chlamydomonas, tz7, 136
Chlamydomonas niualis, pl. tja,
1Jc
Chlorobium uinosum, roi-ro2
clrlorofluorocarbons, ro6
Chloromonas, pl. t jb
chlorophyll, rot, ro2, 2o3
chlorophytes, zo3
chloroplasts, zo3, pl. zo
choice, 5, 7, 164-165,2r7,2r8-22o, corn, 2o8-2o9
222, 223-22+, 226-229, 23r-232, cows, r3, 89, rzo
233
chordates, 16r-16z
Christchurch Press, z3g
Christianiry 5; and science, 7-8, 35,
37-40,45-46,48, 52, 54, zr8,
23r-232; spirit and, 36
Chromatium, 59
Chromatium uinosum, pl. 7
chromosomes: bacteria as lacking,
94; evolution and, 47; of humans, cultural selection, 22r, z3o
r16, r3r;meiosis and, r37, r39,
r4o; mitosis and, r23, rz6; numbers curiosiry, zr4
oi 116; of plants, r97. See also
DNA; genetics; RNA
ctha, tz7, r37,2o3
cities, zo9, z16
clams, t34
(Copeland), rr8-rr9
classification oforganic beings:
kingdoms, r 16-r 19, r4r, r71,
r72, r8r,2r4-2tS phyla, r45,
r47, r5r, r53, t7S-t77,r95;
species (see species). See a/so
animals; bacteria; fungi; plants;
prorocrists
clays,74-75
Cleveland, Lemuel Roscoe, r3g
coal, 176, tg6, rgg, zo6, zro, pl. zj
coal balls, zo6
coccoids, rro
coccolithophorids, t4z, 238
Cohen,Yahuda, ro9
contets, 206-207,24r
conidta, r77
conifers, zo7
conjugarion, 94-96
connectedness. See Gaia
consciousness. See mind
cooperation vs. competition, 23, t6g,
2to,2r+,236, 238. See also
symbiosis
Copeland, Herbert F., r r 8-r r9
Copromonas, zt8
cora-ls, 163, 238
crabs, r48
Craniata, r47, t67
Cretaceous period, 79, r8r, zo8
Crick, Francis, 7, 8, 60
crown gall disease, r8r
cryptozoans (stromatolite$, ro9-rro,
pl. g, pl. toa-b
crystalline structures, r, 27, 7 5
cucumbers, r35
culture, wealth and, zoo
Cuvier, Georges,66
cyanoacerylene , 7 3-7 4
cyanobacteria (blue-green): atmos-
phere and, 89, go, gz, ro3-ro6,
rzg; chloroplasts and, pl. zo; in

desert crusts, 2o2-2o3; evolution
of, ro3-ro4, r95; Fkcherella, pl. 8;
geological effects oi ro5-ro6;
lichens and, rzo, r77-t78; in
microbial mats, r08, ro9-r ro,
zoz; mitochondria and, 9o; plastids
and, rzr, pl. tja;pollutionand,
ro6. See alsobacteria
cycadofilicales, 20 s-zo6
cyclosporine, r88
cysts, r4r, 160,24r
Damiani, Cardinal Pietro, 64
Daptobacter, r3t
Darwin, Charles: Buder's challenge
to, 222-zzg, 239; on complexiry
5, 20l as evolution theorist,45.
47-48, zz3, zz5-zz6; on flowers,
zo8; Haeckel and, 44, 45; on
individualiry r36; on life, origina-
tion of, 69; Lyell arrd, 46;' material-
ism and, 4r; and natural selection,
g, r45, z2o-22r; organic being, as
term, r4; schooling of, zz3
Darwin, Erasmus, 68, zt3, 223, 224,
225
"Darwin among the Machines"
(Cellarius), 239
"Darwin on the Origin of Species,
a Dialogue" (Buder), 239
Deamer, D*id, pl. 4
death: animals and, r56-158; and
autopoiesis, fet\tre of , 7 8*7 g;
bacteria and, 9r, 97, r36-137;
continuity and, zz8; evolution
of, r37' fear of, purpose of, 3o-
3r; funerary rites, 34, t86, zzr;
as great leveler, zoo-zoz; as
illusory, 8r; mystery of , 33-34;
programmed, t37, r44, r53,233;
protoctists and, rr4, r36-t37,
r4+, r56; recycling and, 9I, r99;
recycling, fungi and, 172, t74,
r8r-r82, r86, r89-r9r; selective,
233-23+; sexualiry and, r36-t37,
r5r, r56-158
deoxyribose,83
I ndex I zzs
DeRerum Natura (Lucretius), 54
37-4o, 65, zzo,
Descartes, Ren6, zzr,
231-232
Descent oJ Man (Darwin), 4r
desert crusts, 2o2-2o3
desire: death and, 8o, 8t. See also
choice
desmosomes, r53
dessication, ro8, r4o, 167,169,
2o2-2o3, 2o4, 2o7, 2+r, 242
deuteromycotes, r77
Devonian period, zo5, zo6
Dialogue of the Tbo Chief Woild Systems
(Galileo), 39
diatoms, t4r, r43, t4j, pl. zz
Dickinson, Emiy,245
Diffinbachia, z7
digestion: animals distinguished by,
r54; gastrulation and, I54; spore
dissemination and, r83, r84,
r 8 5-r 86; symbiotic development
and, r33-r34, r3g-t4r, r4+;
unconscious memory and, zzg.
See also food
dimethyl sulfide, r4z
dinomastigotes, r 3 8- r 3 9, z 4o
diploids, r+o, rg7, rg9
disease: bacteria and, 87-88, roo;
death as, r37; fungi and, r8z, r88;
protoctists and, r4r; symbiosis vs.,
r2o, r3r-r32
disequilibrium structures, 9r, 97
disorder. See autopoiesis; entropy;
thermodynamics
dissipative structures/systems, r 6,
57-58, 76-78, 8r, 82, r5r,
pl. 5a-b
divine right of kings, 4o
DNA (deoxyribonucleic acid): bac-
terial, 89, 94-96; of centriole-
kinetosome organelle, rz7, r2g;
discovery of, 7, 8; of eukaryotes,
r14, r16, rz3; evolution and,47,
86; function of, 8z; mitochon-
drial, r33; plastids and, rzr;
production of. 29, 83; proteins
and production of, 58; and RNA,
 276 lndex
DNA (continuetl)
8z-83, 85; synthesis of nucleic
acids, 73. See a/so chrornosomes;
genetics; RNA
Dobzhansky, Theodosius, 47
dogs, r48, rJr, r68
dolphins, r48
drugs: fungi and, r83, t86-r87, rgr,
zr5,238; plants and, r94, pl. z5
dualism, 48
dulse, r 3 5
dung, r83-r84
Dynastes, t 46
Dyson, Freeman, 75
Earth: as alive, xiv, 23-24,46,52-55,
r68-170, r89, r90, z37,pl. ta,
pl. z6; atrnosphere of (see atmo-
sphere); biosphere of (see bio-
sphere); formation of, 6o-6r;
asholarchy vs. hierarchy, 9, 9o;
noosphere of, 4g-5o, r7o, 23 3;
rhythms of. z4o-24t. 242i rimc-
line of history 6z-8o; viewed
fromspace, ro-r2, r68, r89,
pl. ta, pl. 26. See also global
environment
ecology, as term, 44
economy: of solar energy, r9o-r9r,
r 99-202, 2ro-2 12, 21 5, 23 4-23 5,
242, 246
Edelman, Gerald, 233
Ediacaran fossils, r6z
eggs: evolutionary asymrnetry and,
rJ8-r j9; fertilization of (see fertil-
ization); immortality of, r56; of
plants, r96, r97,pl. zq
Ehrenberg, Christian Gottfried, 46,
47
Eigen, Manfred, 83-84
elements: as stellar rnateia\, 25, 3z
Eleusinian mysteries, r86
Ellington, A. D., 84
emboitement, 66
embryos, r S2-r 53, 234; blastula, r45,
t46, r5z-r54; plant, r5z-r53,
196-199, 198, pl. 24
Emerson, Ralph Waldo, r4_5
Emiliana huxleyi, 5 j
endorphins, 30, rg4
endosymbiosis. Scc symbiosis
energy. See food; fossil fuels; metabo-
lism; solar energy
England, 4t,45-46, r88
enkaphalins, 3o
entropy, r5-r6; autopoiesis vs., 43,
8o-8r, 86; life and, r5-16, t7, rg.
See aLro thermodynarnics
environment. See Earth; global
environment
Epilagus, t96
Ereuhon (Butler), zz3
Erythrodinium, r66
Esclrcrischia coli, 96
Eschinkcus blumi, t69
Eucharist, r87
Euglena, t26, t35
eukaryotes, r14, 1 15, r r6; discovery
of, xii-xiii; symbiosis and (see
symbiosis)
Europe: Cartesian license and, 4o-4r,
45-46; fungi and, r87-r88; and
male life essence, 65; medieval, 3,
36-37, +o, 17r
European Space Agencies, z4z
evolution: abuses of theory. 48; of
anrmals, rJr-r52, r54-r56, r59-
t64, 167-17o, zo7-2o8, zog,
p/. 15; autopoiesis and, 8r; of
bacteria, go,93, g7-gg, 154;
Butler's challenge to Darwinian,
222-229, 239; Cambrian vs.
Precambrian, r 59-164; Darwin
and, 45, 47-48; of death, r37;
deception and, r65; definition of,
r8-r9; ofeyes, r65-167; offungi,
r76, r77-r8t, 196; of humans,
r5z, i58-r59, zo7-2o8, zog,
z16-218, 222, 223-224, 234-2+o,
z4t 243; mechanism and,zz3,
225,23r-232; mutation and, 8,
rzr, tz8; novelry in, source of,
8-9; opposition to theory oi 45-
46,66; of plants, r79-r8r, r94,

ry 5- t 96, 202-2 t o; in population
of one, 23. 53i pre-Daruinian
theories and,45-47; as process vs.
mechanistic law, 164-167, zz3; of
protoctists, 27, 93, 94, r t6-r 17,
r 54, I 59-I60; purposefulness
and, zz t*zzz, 22 3-22 +, 226-2 3 r,
z3 5; symbiosis and (see symbiosis);
technology and, z16-218, 238-
2+o, 2+r-2+3; of wood, r76,
I88-r89, zo6. See a/so autopoiesis;
reproduction
Evolution, Old antl Neru @ut1er),
224-225
Expression oJ Emotions (Darwin), 4r
extinction: end-of-Cretaceous, zo6;
fossil record and, zo6; modern,
2rr, 245-246; of Permo-Tiiassic,
206-207: solar economy and,
r9o*r9r, 2or-2o2, 245-246;
species longevitl and, zt5-2t6.
See a/so deathl populations; species
eyes and vision, ro3, rz7, r65-t67
Faasr (Goethe), 43
Gar, r65; and disdain ofbacteria, 58,
59, 88, 9o-9r, rr8, zrg-zzo
females: evolutionary asymmetry and,
r 58-r 59; sexual choices of, 164.
See a/so sex
fermentation, 99-roo, ror, r88
Fernel, Jean Franqois, 36
ferns, rg7, 2o3,2o5
Grtilization, 47, r45, t46, t5z, r58;
of plants, 196-197, 2o3, 2a4, 2o7,
pl. 24. See a/so sex
fertilizer, ro7
Fkcherella, pl. 8
fish, r58, pl. t8a-b
flagel7a,9z
flamingos, r34
flarworms, t2o,166
Fleming, Alexander, r88
flowering plants, r95, 2o7-2o9, 2r S,
pl. 24, pl. z5
Folsome, Clair, zI6
food: agriculture, r3j, r84-r85, zo8-
lndex 277
2og, zro, zr r ; animal behavior
and procurement of, r5o-r5r;
bacteria and,92, gg-roo, t23,
zr8; cannibalism, r3g-r4r, t44:
essential nutrients and, 8z; fungi
as, r83-r86, r87-r88; fungi, and
procurement o{, t7z, r8r-I82,
r84; heterotrophs and, roo-ror;
laboratory origination of, 7z; per-
ception of, and survival,27, 3o;
plants and, 196 (see a/so photosyn-
thesis); protoctists and, I 33-r 34,
zr8, zzo, zz4; and sex, origins of,
r 3 9-r 4 r ; storage of energy and,
r99; sugars (see sugars); symbiotic
development and, r 33-r34, r39-
t4t, t+4. See also digestion
foraminifera, t+r, r43,2r9, 48
forests, r76, I88-I89, 206,2rr
fosil fuels, igg,2o2i 2ro-zrt, zrs;
coal, ry6, t96, rgg, zo6, zro,
p|.23
fossils,zz, 61,66,82; of animals, r47,
t5z, r59-164, 168, zo6; ofbac-
teria, 6r, 88-8q, q8, to7, ro9,
t ro, pl. t oa-b; fvngr and, zo3;
of plants, 195,2o3, zo4, zo6, zo7,
zo8, pl.2J; protoctists and, r59
Fox, Sidney, 7I
fractals, 4, go, 174,2r7,46,242
France, r88
free radicals, ro4
free will, 5,2r7,222,23r-232. See
a/so choice
French Revolution, 4o, 45-46
Freud, Sigmund, zzr
Frisch, Karl von, r49
frogs, r58, 166
frutfly, pl. t6
fruits, r8o-r8r, 2o7, zo9
Fuller, R. Buckminster, 8
fungi (Mychott), 173, pl, t9b;
animals and, coevolution of, r83-
r86; animals as preceding, r47;
attitudes toward, r18, 17r-t72,
r75, r 83, r86-188. I89, I90; in
desert crusts, zo3; evolution of,
 27A lndex

fungi (Mychota) (continued) Glaessner, Martin, r6z

176, r77-r8r, r96; as food, r83- globai environment: as closed system,
r86, r87-r88; food procurement r9o-r9r, 2or-2o2, 24r-243;
^s
by, 17z, r8r-t82, r84; function interwoven, tg-zo, z3-24; and
of, 17z, 174, 176, r8r-r82, r88- minerals, life as, 49; mrrror stage
r9r; individualiry and, 174-r75, and, r r; popuiations regulated by
r 89; as kingdom of liG, rr 8-r r 9, solar economy in, 98-99, r9o-
r7r, r7z, r8r, zr4-zr5; as land r9r, 2or-2o2, 234-235, 246;
beings, r7z, r74; mineral use and success of life forms and, r 89,
production by, z8-zg, r7g, r8t I 90, r gg-2o2i superhumanity
mitosis and, t23, r26, rz8; phyla a.nd, 234-235, z4r-243; rse of
of, 175-177; plants and, r78-r8r, matter by life and, 27. See also
196,2o3; reproduction of, r7z, Earth; extinction
123, t74-r75, t76-177, r8z, Clossopteris, zo6; C. scutum, pl. z j
r83-r86; symbiosis and, ry7-r8r, glucose, gg,232-233
pl. tga; as term, r9o glycine, Tz
gnosticism, 3 6-3 7
Gaia, xiv, +6, 52-55, r68-17o, r89, God, 5-7, 36-37, 38, 4o-4r, 54,66,
rgo, pl. ta, pl. z6 22r,23r-232, pl. + kap.)
Galileo Galilei, 39 Godel, Kurt, 4r-42
galls, r8o-r8r, r8o Goethe, Johann Wolfgang vorr, +3-44
gtmetangta, t76 Golubic, Stjepko, r95
gametes. see eggs; sperm Gondwanaland, zo6
gapjunctions, r53 gorillas, r48
gastrulation, r 53-r j4 Gould, StephenJay, rr3, r52,22t
gender: bacteria and, 94-96; evolu- Grapaldus, Franciscus Marius, r7r
tionary asymmetry and, r58-r59; grass-green bacteria, r 34
fungi and, 176. See also sex graviry 6, 7, ro, rt, +g
genetic code, 82 Great Chain of Being, 9, 4r
genetics: bacterial gene trading, 93- Greece, ancient, 35, 42, 59,64, 186
96,95,97, rrr, r16; evolution greed, zoo
asexplanation of, 47; human greenhouse gases, 22, 24, 2o2,2rr
evolution and, z16; as window to green perspective, z4z, 243
evolution, 98, t2t, r3o-r3 r. See green sulfur bacteria, ror-r02, r07,
a/so chromosomes; DNA; RNA I o8-r 09
gene trading, 136 Griffin, Donald, r49-r Jo
genitals, r58 gulls, r48
geology, 45-46, 49,5o, 88-89, r67; gypsum, 2Io
cyanobacteria and formations
of, ro5-ro6; plate tectonics,206, Hadean eon, 6r, 65, 6+-6 S
pl. z 3, pl. z 6. See also fosstls; Haeckel, Ernst, 44, 45, 48, 68-69,
minerals rr7-rr8
Germany, r88 Haemanthus, 6tz
germ theory 88 hair, 165
Gilbert, Walter, 85 Haldane, J. B. S., 7r
Cinkgo, zo3, zo5 Ha1l, John, r27, r57
glaciers, 206-207 Hallucinogenia, t6t

hallucinogenic mushrooms, r 83, r86-
r87, r9r, zr5,238
Halobacter, t66
halophiles, 89, ro2-r03, I66
haploids, r4o, r96-197, rg9, pl. z4
Harvey, William, 6, zz9
HCN,74
heat, r 5-r7 . See also entropy; solar
energy; thermodynamics
Heisenberg, Werner, 4r
HeLa cells, I35
helium, z5
Helmont, Jean Baptiste Yan, 57,
6+-6s
hematite, ro5
hemoglobin, g8
Hepburn, Katharine, z r7
herons, r48
heterocysts, Io7
heterotrophs, roo-ro3, r 54
hierarchy, holarchy vs.,9, 90
Hinduism,48, I86,2I8
"Historical Sketch of the Progress
of Opinion on the Origin of
Species" (Darwin), zz3
Hogg,John, ir7
holarchy, 9, 9o
holons, 9
Homo sapiens. See humans
hornworts, zoz
Horodyski, Robert, r95
horsetails (scouring rushes), 27, zo5
humans: aging of, gr, ro4; as animals,
xi, xiv, r49-r5o, r5z, zt5-216,
2r7; attitude of uniqueness, r7,
22r, 224, 23o, 246; cell replace-
ment in, I7; chromosomes of,
r16, I3t; as consumers in solar
economy, r90-r9r, r99-2o2,
2 ro-21 2, 23 4-23 5, 245-246;
deception and selGdeception in.
t65, zzr; descendants of, evolu-
tionary zr6; essential amino acids,
8z; evolution of, t5z, r58-r59,
2o7-2o8, zog, zt6-2r8, zzz,
22 3-224. 2 3 4-240, 2 4 r-2 43 ;
fungi and, r83, r85-r88, r9o-
lndex 279
19r; gender asymmetry and,
r58-r59; gooseflesh, r65; hair-
sryles of, r65; intelligence of, r65,
212,216, zr8; mirror stage of, rr;
as planetary proprioceptor, r68;
plant-human interdependence,
zo8-zog, 2to-2r2, pl. z 5; as prey,
9o-9r; shelter and, z4r-243; stress
metabolism of, 85; superhumaniry
zt7-z 18, 23 4-240, 24r-243;
survival of, xiv-xv, 2+2-243,245-
246; symbiosis within, rzo, 236-
237; technology of (see technol-
ogy)l timeline of, 6z-63; vision
of, ro3, r27, r66-t67. See dlso
animals
Humboldt, Friedrich Wilhelm
A-lexander von, 46-47
Humphries, Nicholas, xi
Hutton, James, 45, 46
Huxley, Thomas Henry, r 18, 169, 226
hydras, r34
hydrogen, 25, 6o, 73, 8r, roz, ro3,
ro4
hydrogen cyanide, T4
hydrogen sulfide, roz, ro3
hypermastigotes, r 39-r40
hyphae, 174, t76, r77
identiry 19. See also individualiry
incompleteness theorem, 4r-42
independence, as term, 2o
individuality: animal embryos and,
r53, rJ4; continuiry and, zz8;
fungi and, r74-r75, t8g; as
illusory, 48; plants and, r54; pro-
toctists and, rr4, rJ-3; as relative,
r36; and symbiosis, r20,236-237.
See also artopoiesis; mind
infrared radiation, ror
insects, r7z, r83
instinct, 2 I 9
intelligence: human, t6S, 212, 216,
zr8. See also rrrind
kaq, 34
iron,28, r05,2ro
Iroquois,34
 280 lndex

Islam, 5, 37,218 Life, r87

isobutyric acid, r86 life: age of, 88-89,97; body of,4o5;
Italy, r88 chemical composition of, z4-27,
3 r-32, 5o, 5r-52,73,8o-82;

Jacob, Franqois, 57 definitions, as transcendant of,

'William, r
James, 3 2I5; entropy and, r4-16, 17,
Jantsch, Erich, 77, 79 I9; evolution of (see evolution);
Janus, 9 fractionation point of, 58; king-
Jeon, Kwang, r3r-r32 doms of. I r6-l r9; as living
Journal of the Proceedings oJ the Linnean matter, 49-52; matter use by,
Soriety oJ London, Zoology, 45 27, 5z; mind as universal to (see
Joyce, Gerald, 84 mind); minimal unit of, r8, 58-
Judaism, 5, 36, 2r8 59. 78, 82. 85-86. r 55; origins
Jupiter, 37,6o,74, ro4 of, 57-6o, 61, 69-7 5, 77-80,
82, 83-86, ro4; purposefulness
Kant, Immanuel,4z and (see purposefulness); reproduc-
Karo Battak, 34 tion of (see reproduction); self-
kelp, r35 maintenance of (see autopoiesis);
Kendrick, Bryce, r75 spontaneous generation of, a4-ag,
Kepler, Johannes, 3, 6 t t7; survival of, xiv-xv, 32,242-
kinetosomes, t26-r27, r29, r57 243, 245-246; what it is, z-3 ,
kingdoms, rI6-Ir9, r4t, 17r, r72, 3r-32, 55, 86, 9o, rro-rrr, r44,
r8r, 214-zr S. See also animals; I7O, 1gI,2Ir-212, zr3-zr5; will
bacteria: fungi: plants: protoctists to,8o
Klebsorbmidium, z,o3 iignin, r76, r96, zo8
Knauth, L. Paul, r95 Lima scabra, pl. t 5
Knoll, Andrew, 88 Linnaeus, Carolus, 66
Koch, Robert, 87-88 lipids, 57, 75, 85, 86, to4, pl. 4
Koestler, Arthur, g liverworts, 2o2, 2o4
Kooning, Willem de, 3 r Lovelock, James E., r, r2-r4, 49,
Korea, r 87 52-5 5
Koshland, Daniel, zrg Lowenstam, Heinz, 25, z7
Kosnos (Humb oldt), 46- 47 Luck, David, t27, r57
Krumbein, Wolfgang, 54 Lucretius,54
"Lucubratio Ebria" (Butler), 239-240
laboulbenomycetes, r 72 Luddites, 239-240
Lacan, Jacques, r r luminescence, r84, z4o, pl. t 8a-b
Lamarck, Jean-Baptiste, 2 2 4-22 5 Lyell, Charles, 45, 46
language, ry; fungi and, r83; written,
I86-r82, r89 machines. See technology
Laplace, Pierre-Simon de, 6 Magic Mountain, The (Mann), z-3
Laurasia, zo6 magnetite, 28, r05
I-aws (Plato), 3 magnetotactic bacteria, 59, 2rg, 22 o
Leeuwenhoek, Antoni van, 34,65, Malay, 34
87, rr3 males: evolutionary asymmetry and,
leishmaniasis, r4r Ij8-Ij9. See also sex
Le Roy, Edouard, 5o mammals: acquired traits and, zzz;
lichens, tzo, t77-r78, r8g brain development, r56; evolution

of, zo7-2o8,2o9; fungi and,172,
r83-r84, r85-r86; sex cells of,
I j6, I j8; species longeviry of, zI 5-
216. See a/so animals; humans
Mandibulata, r4S, r47
manganese,28, ro5
Mann, Thomas, zo1
Maori,35
Mariner, Ruth, 7r
Mars, Iz, 20,21,90, to4
mastigotes, r39-r4o, zI 8
materialism, 4r, 4+, zt7
Maturana, Humberto, r7
mechanics. See science
meiosis: evolution of, t38-r4o; fungi
and, 175; mitosis and, r37, r38;
plants and, 196, tg7; process of,
r37. See also reproduction; sex
memory. 27. zt St unconscious, zz t-
222, 223-224, 226-229
Mercury, 37
Mesodinium rubrum, pl. tt
Mesozoic era, 79
metabolism: aerobic, 85, r3o, r33;
anaerobic, 85, 89, ror-Io3, rz8,
p/. 7; atmosphere and, 89; auto-
poiesis and, r7-r8, tg,77-79;
components of, 78; definitions of,
r9; nonliving systems and,76-77;
photosynthesis (see photosynthe-
sis); symbiosis and, tzo; as unceas-
ing, 79; as window to past, 5 8-59,
8o-82, 83, 85, 86,98-99, toz.
See a/so autopoiesis; food
metallurgy, z3 8
metals, 27, j2, I05
metamorphosis, r 54, 228
metaphysics, 40, 42,72
meteors, 6o-6t, zo6-2o7, z4r
methane, r 3, 7o, 73, 89, 92, 167
methanogens, 89, pl. z
microbes: defined, r t6. See alsobac-
teria; fungi; protoctists; yeasts
microbial mats, 89, roj, Io9-r ro,
zoz, pl. 9, pl. toa-b
Miootystis, 59
rnicroscope, 34, 65
microtubules, rz3, zo3, pl. t 7
lndex 241
mildew, r8z
Miller, Stanley L., 57, 7 r, 7z-7 3
Mills, Donald, 84
Milton, Jeffrey, r94
mind: animals and, r5o, r5z, 167-
r7o,2r+; autopoiesis and,3t, zzz;
bacteria and, zr8-zzo; cells and,
ztg-22o, zz8-zzg; evolution of,
r64; instinct end, z19; knowledge
of self and others, xi; materialism
and, 4r; and matter, as one, 43-
44, +8-55,233; end matter, as
separate, 37-4o; and meaning of
life, 3; natural selection and, r64,
165, 230-23r, 233-234; noo-
sphere and, 4g-5o, 17o,233
mind, as universal to life, 27, 3o-3 r,
48, r48-r5r, r67-t7o; chemical
and physical processes of, z3z-
234; choice and, 5,7, r6+-165,
zt7, zr8-zzo, 222, 223-224,
226-229, 23 r-232, 23 3 ; purpose-
fulness and (see purposefirlness);
unconscious, 30, z2r-222, 223-
224,226-229, 23+
minerals: life as, 49, 5o; liG as pro-
ducing, z5-29,26, ro2, r43, 163-
t64, r66-t67, 176, r8r, zo5,
237-238; oxidized, and atmos-
phere, ro5-ro6, tzg, 16o
mitochondria: dependence of, I 3 r;
DNA of, r33;respiration and,
r3o; spirochete symbiosis as
preceding, Iz8l symbiosis as
originating, go, r2r, I3o-r3r,
t 32-t 3 3 ; unconscious memory
and, zz8-zz9
mitosis: anima.ls and, r 5 3. r 57; in
blastula, r53; defined,94; evolu-
tion of, 94, r26-t29; plants and,
ry7, pl. tz; Process of, r23, 125,
rz6; telophase, pl. tz. See also
autopoiesis; reproduction
mitotic spindle, r23, 126
Mixotricha parudoxa, r z3
mo1ds, r75, 176, 177, r88. See a/so
fungi
Monera. See bacteria
 282 lndex

money,2oo negentropy, r5-16

monism,44 nematodes, r84
monkeys, r48, r65 Nephromyces, z6
Monotropa, 196 Neptune,6o
moon, Io, 71,72,37,6I nervous system, r j3, r56-157,23o,
morality, 4o 233-234
Morison, Robert, r Newton, Isaac, 6, 45, 48,65
Morowitz, Harold, 85 New Zealand, 3 5
mosses, 2o2, 2o4 Nietzsche, Friedrich, 4r
motility: bacterial locomotion, 92, Nigeria, 234
ro2, 122-123; symbiosis of, rz3, Nightingale, Florence, 88
t26-129; undulipodia, 124, 126- nitrates, ro7
tz7, tSS, r57, 166, zo3, pl. t7 nitrites, ro7
multicellularity: as bacterial invention, nitrogen, 13, ro7, r8z
9o, pl. 6; complex beings and, nitrous oxide, 13, to7
r3 5-r36 noosphere, 4g-So, r7o, 233
muscles: blastula and, r53 Nostoc, 59
mushrooms, ry6-r77, r8 j-r86, notochord, r6r
r 87-r 88; hallucinogenic, r 83, nuclei, 94, rr4, 1 15, r16, tzg, 13S
I86-187, rgr,2rS,238. See also nucleic acids, 7 3, 78, 85
fungi nucleotides,83
mutation: evolution and, 8, rzr, rz8 Nymphaea, ztr
Mutinus, r83
mycelial network, t7 4-r7 s oak trees, r98
Mychota. See fungi oceans: animal evolution and,
mycoplasms, 82, roo,pl. tb r 5 r-r j2; autopoiesis and, zz-23;
mycorrhizae, r78-t7g, t9g fungi in, rarc, r72, 174;
^s
Myxococcus, pl. 6 protoctists and, r4r, t42-1+3,
rg4. water
See also
Nabokov, Vladimir, t64, t65 "On the Distinctions of a Plant and
Naegleria, t j I an Animal, and on a Fourth
NASA (National Aeronaurics and Kingdom of Nature" (Hogg), r r7
Space Administration), 12, z4z On the Origin of Species (Darwin):
native Americans, 9, 34, 3J publication of, 4r, 47-48
Natural Hktory @uflon), 68 or)togerly,227
natural selection: artificial selection Opabinia, fir
vs., z2o-22r; cultural selection vs., Oparin, Alexander Ivanovich, 69-7o
zz r; function of, 9, zot-zoz; orangutans, r48
materialism and, 4t; rnind and, Ordovician period,, 63, Z 7, 2o2-2o3
164, t65, 230-231, 233-234; organelles: definition of, t2t;
survival of the fittest, tg, 48, 53 evolution of, t2r, 126-129, r32.
Nauicula cuspidata, pl. z z See also cbloroplasts; kinetosomes;
Nazis, 88 mitochondria; plastids; symbiosis;
Neanderthal rnata, 34 undulipodia
Neckam, Alexander, 64 organic being, as term, 14
necrobes, r3r Orgel, Leslie, 57, 7r
Needham, John Ruberville, 68 Origin oJ LiJe, The (Oparin),7o
 lndex 243

Origin oJ Species (Darwin), t 4, zo, 196; on land, development of, r95;

lichens and, r77, t781' microbial
ownership, rgt, 2oo-2or mats and, ro8-r ro; pigments and
oxygen: biological production of, r3, oxygen tolerance in, roz-ro3,
89,9o, ro3-ro5, r06; and pl. 7; process of. ror: variations
deoxyribose, 83; methane of, gz. See also metabolism
production vs., r3; origins of life phyla, r45, t+7, rfl, r53, t75-177,
and,7t; respiration of, rzr, r3o, r95
r 3 t, z z8- z z9 ; toxicity/tolerance phylogeny, zz7
of, 59, ro3, ro4, ro6, rz8, pigments, r2r, t34-r35, 166,2o3,
r2g-r3o. See a/so atmosphere pl. tja-c; oxygen tolerance and,
oysters, I 5 8 toz-to3, pl. 7
ozone layer, ro6 pigs, r85
Pikaia, r6t-r6z
Pachnoda, pl. z Pilobus oystallinr.rs, I 83- r 84
paleontology, r59 pineal gland, 39
Paleozoic era,6j, 77-78, 176 Pirozynski, Kris, r79, r80, r8r
Panisset, Maurice, 87 Plachobranchus, r34
panspermia, 59-6o plants: algae as ancestors to, r7g,
Papauer somniJerum, pl. z 5 ry+, ry5-rg6, zo3; animal-plant
Papua, 34 interdependence, 2oo-zor, 2o7-
Paracoccus denitrijcans, r3 r zro; animal-plant split, t18, r97,
Paramecium, I35, zI8 r99; animals as preceding, r47;
parasitism, r3r, 216. See also symbiosis bacteria and, 93, 97, pl. tja,
Parmenides, zt3 pl. zo; embryos of, r5z-r53,
Pascal, Blaise, zI3 196-199, tg8, pl. z4; evolution
Pasteur, Louis, 69, 87, 88 of, r79-r8r, ry4, t95-r96,
pattern recognition, 3o 2o2-2ro: flowering, tg1, 2o7-
Pedomicrobium, 238 2og,2rS, pl. 24, pl. z5; function
Peirce, Charles, 3 r of, r94-195, zrr; fungi and,
Pelomyxa palustris, r 38 r78-r 8 r, ry6, 2o3; human-plant
penicillin, r82, r88 interdependen ce, 2o8-2o9, 2t o-
peristalsis, zz9 zrz, pl. z5; individuality and, r 54;

peroxidases, rzg as kingdom, II8, r4r,2r5; as

Phallus, 183 landbeinp, t47, r7g-t8o, r9+;
Phanerozoic eon, 6 z-6 j, 7 7-8 o, 16o, mineral use and production by,
zr8 28-29, zo 5, 237-23 8; mitosis
pharmakon, 186 and, rz3, rz6, rz8; modern
Phormidium, rz9 descendants of earliest, 2o2-zos;
phospholipids, p/. 4 multicellulariry of, r 3 5-r 36;
phosphorus, r8r, I8z plastids, 90, roo, rzr,133-r35,
photosynthesis: atmosphere and, 9o; as r66; protoctists as ancestors of,
basisof liG,23, Jr, ror; economy t16-117, r33, r4r, r94; reproduc-
of globe and, r99-zoo, 2or, zoz, tion of, 94, tg6-rgg, 198,2o3,
zro-2r2, zr5; evolution of, ror- zo7-2ro; seeds of, zo5, zo7, zo8-
r03, r9j, pl. z, pl.zo; hydrogen zog,2+t; and sex, t96-r99, pl.
sulfide and, roz; lack of, in plants, z4; size of, r94,2o4, 2oj; spores
 284 lndex

plants (continued) protoctists, pl. t t; anddeath, I r4,

of, I9J, t96-t97, zo4;waterand, r 36-1 37 , r 44, r 56; and disease,
r95-196, 2o2-2o3, 2o4 t4r ; evolution of, z7 , 93, 94,
plasmids, 94, 9j, rr r, r8r r16-1t7, r54, rj9-r60; function
plasmodesmata, zo3 of, r4t-r44; individualiry and,
plastics, 27, 52, ro8 rr4, rj3; as kingdonr, rr6-rr9,
plastids, go, roo, tzt, t33-r35, 166 r4r; light sensing and, r66;
Plato,3,36, r86 nreiosis a'nd, r37-r4o mind and,
pleasure, survival and, 3o zt8-zzo, zz4; mineral use and
pollution: humans and, ro6, i64, production by,25, 26,28-29, r43,
rBz,245-246; as natural process, 237-238; mitosis and, r23 r2g,
ro6-ro8, rrr, 163-164, 167; ,25; mutation and, r.z 8; ocean
plants as purifying, zr r; popula- and, r4r, 142-r+3, r94; as pro-
tion growth and, 99; recycling genitors, r33, r4r, r58, r6o;
(see recycling); solar econonry recognition of one anothcr, r4o
and, zro-.zr t. See also autopoiesis; r4r; reproduction of, r37-r4t,
thermodynamics r56, pl. 22; sex and, r36, r44;
polyaminomale onitryl, 7 4 symbiosis and (see symbiosis)
polyps, r55 protonritochondria, r 3 r
Ponnanrperuma, Cyril, 7 t-7 2, 7 4, Proton motors, 92
pl.s kap.) protozoa, as term, r I7
populations, regulation oe 98-99, Psilocybes mexicana, r87
r 90-r 9 r, 20 r*2o2, 23 4-23 5, Psilotun nudum, zo4, zo5
246 Pteridinium, r6z
Populus tremuloides, t94, pl. z t purplc bacteria, g5, ro3, roj, r09,
Porphyridium, tzr t3o t3r, pl. 7
Prigogine, Ilya, r6 purposeFulncss (teleology) : auropoicsis
primates, zo7-2o8, zo9 and. r(r 17. 52-Jl; Jefinirion
Principles oJ Ceology, The (Lyell), a6 of, r7; ordinariness of, 224,23r;
Prochloron, t34 science and, 17, 224, 227; thermo-
Prochlorothrix, r34 dynamics and, 16-17,86, r5r;
prokaryotes, 94, rr4, rr5, rr6. See unconscious memory and, z,zt-
also bacteria 222, 223-224, 226-231, 235
propagules, z4r pyrite, to5
proprioception, r 67- r 68
proteinoid mrcrospheres, 7r Qtrercus, 180, 198
proteins: autopoiesis and, r9; histone,
r 3 r; origin of liG and, 69, 7 r-7 3, radiolaria, r4r, t+3
7J; production oi S8, 82, 8:, 85; recycling: closed system of biosphere
reproduction and, 79, 8o and, r9o-r9r, 2or-2o2, 24r-2+3;
proteobacteria, g5, ro3, roj, r09, fungi and, rllr-r82, r89-r9r;
t3o-r3r, pl. 7 humans and, t8z; as natural
Proterozoic eon, Z1-ZZ,93, 94, rr1, process, ro6-ro8, ttt, t63-t64,
r44, r56, r60, rgJ I67; respiration as, r3o; savanna
Proteus mirabilis,pl. 5a evolution and, zo8. See alsoEarth;
protists: as term, r17. See also pollution
protoctists Redi, Francesco, 67

red pigment, ro2-ro3, r2r, r34-r3 5,
166, pl. t t, pl. tja-r
religion: animism, 3, S,8, 17, 33, 4r,
zr7; censorship through, 37, 39;
and common heritage of liG, +8;
death and, 34-37,22r; fungi and,
r 86, r 87; origin of life and, 66,
69; and science, 7-8, 35, 37-40,
45-+6, 48, 52, 54, zr8,23r-232;
and sun, importance of, 5r. See
also atheism; metaphysics; spirit
replication, vs. reproduction, I8
reproduction: of animals, 94, rSr,
rjJ, r58; autopoiesis as preceding,
r8, 43, 79-8o; of bacteria, 79*8o,
91, gz, 94 96, 95; of biosphere,
242; economy of, r89, r9o; of
fungi, r7z, t7j, t74-t75, 176-
r77,182, r83-r86; meiosis (see
meiosis); mitosis (see mitosi$; par-
thenogenesis, 66; of plants, g 4, 196-
t gg, t g 8. 20 3, 2o7- z t ot of pro-
teins,79, 8o; of protoctists, r37-r4r,
r56, pl. zz; sex (see sex); vertical
vs. horizontal, 94; viruses and,
94, 95, 96.Scc a/so autopoiesis:
evolution
reptiles: reproducrion of, t58
respiration, r2r, r30, r3t, zz8-zz9
retinal, r66
Rhiuzopus stolonifer, 17 6
rhodopsin, ro2*r03, r2t, r34-r35,
t66,pl. tt,pl. rja-c
Rhynia, zo3, zo5
ribose, 83,99
ribosomes, 83, 89
ribozymes, 84
Rig-Veda, t86
RNA (ribonucleic acid): autopoiesis
and,79; bacterial, 89; of centriole-
kinetosome organelle, rz7; DNA
and, 8z-83, 85; evolution and,47;
function oIl 8z-83, 84, 86; origins
of life and supermolecule of, 75,
8z-85; replication of, 83-84. See
also chromosomes; DNA; genetics
Robinson, Jennifer, r96
lndex 285
Roman Catholicism, r87
Rome, ancient, 37
Rosenbaum, Joel, tz7
Russia, r 87
Russula paludosa, pl. t ga
rust, Io5, I60
Sade, Donatien Alphonse Franqois de,
40
Sagan, Carl, 7r, 74
Salix, t94
salts: desalinization of, zz-23, 52. See
a/so minerals
Saturn,37,6o,74
savannas, zo8
Schizophyllum comrrune, pl. t9b
Schrodinger, Erwin, r-2, z, 7, 8, +7,
71,76,245,246
Schwartz, Karlene, rrg
science: as asymptoric. 3; autopoiesis
and, j3; Cartesian license of, 37-
40, 52, 54,22r,223, z3z; and
context, 42; contrary evidence
and, 67 -68 ; departmentalization
of life and, r 18; future of, zr8;
intuition and, jo; limitations
of mechanistic,7-8, 53, 54, 55,
2zg-23o: materia.lism and, 4r, 44,
zr7; rnd mind-matter split, 3 7-
4o; origins oflife and,7r; religion
and, 7-8, 35, 37-+o, 45-46, 48,
52, 54, zr8,23r-232; social effects
of, 4o-42; and spirit, 35; teleology
tnd, 17, 224, 227; truth and, 3o-
3r; uncertainry and,4z
sclerites, r6o
scouring rushes, 27, zo5
sea slugs, r66
seasons: planetary breath and, z3-24
sea urchins, r58
seaweeds, rrg, t2r, t3S
Sedgwick, Adam, r59
seed ferns, 205-206, zo7, pl. zj
seeds, zo5, zo7, zo8-zog, z4r
self, animal recognition of, r48
self-determrnarion. See choice
self-maintenance. See autopoiesis
 285 lndex

sentience. See mind tion(see extinction); fixed, 65-66;

septatejunctions, r53 fungi, number of, 175; Iichens,
sex: bacteria and,94*96; death and, number of, r77; longeviry of,
t36-r37, rjr, rJ6-rj8; evolution zr 5-216, 236; plant, number of,
of, t37*r4t; food and, r3g-r4t: r95; protoctists as forming, r36.
fungi and, r72, rBS; gender asym- See a/so of organic
classification
metry in attitudes toward, r 58- beings
rJ9; plants and, 196-199, pl. z4; sperm: evolutionary asymmetry and,
protoctists and, r36, t44. See ako r58-rj9; fertilization and (see
reproduction fertilization); homunculus and, 34;
Shakespeare, William, 8r, r45 immortaliry and, r56; of plants,
Shanidar ceve, 34 196, rg7, pl. z4; undulryodia of,
sharks, r68 rz6
shells, 25, 27, 163-164,2r9,237 sphagnum moss, 2o4
silica, r4, 25,27, 28, r09, r ro, t+3,20s Spiculosiphon, zr9
Silurian period,6j, ZZ, r7g-r8o spider plants, zr r
skeletons, 25, 27, 163-164, 237 Spiegelman, Sol, 83
sleeping sickness, r4r Spinoza, Baruch, 8, 44, 54
slime rnolds, r37, r4r spirilla, r ro
Smith, David C., rz7 spirit, 5, 33,34-37,44, zr8. See also
snails, I34 mind
snakes, r68 spirochetes, rro, rz2, r23,124,
Socrates, r86 r27-r29
soil, r9-zo, r77, r8r, r89 Spirulina, 59
solar energy: bacteriophage produc- sponges, r55, rJ8
tion and, 96; as basic to liG, 5, spores, r j9. r 90, 2+t: of bacteria.
43, Sr, 53, r94; metabolism and 59, 87, rro, rSg,24r; offungi,
(see photosynthesis); surplus and \72, r74, r83-r84, i85-r86, r89,
economy of, r9o-r9r, rg9-2o2, r9o; human shelter as, 24r-243;
2ro-2r2, 2t5, 234-235, 242, 246. panspermia theory and, 59-6o;
See a/so sun of plants, rys, 196-197 , 2o4; o{
solar system: early beliefi and, 3, 36, protoctists (cysts), l4i, 160, 2+r.
37: life potential elsewhere in. lz, See a/so reproduction; seeds
ro4 staphyla, t85
solipsism, zzo stromatolites, to9-tro, pl. 9,
Sonea, Sorin, 87, 136 pl. t oa-b
soul, 34-4o, 44 strontium,28
sound waves, 73 Suess, Edward, 5o
SovietlJnion, ro,7o sugars, 85,99-Ioo; ATP and storage
space: compounds of life present in, of, ror, r3o; deoxyribose,83;
25. 32. 73-74; spore transmission fruits, r8o-r8r, zo7, zog; rlbose,
through, 59-6o 83, 99. See also food
space exploration, Io-r2, r4, r3o, sulfur, roz, r4z
r3J, r68, zrr, zt6,23g-z4o sulfur bacteria, tor-ro2, ro7,
Spain, i87, r88 r o8-r 09
Spallanzani, Lazzaro,68, r17, r47 Sumatra, 34
species: animal, nunrber of. r5 r; su-n,22, 32, 37, Sr,6o, pl. j; energy
bacteria as single, 93, r36; extinc- of (-see solar energy)
 lndex 247

superhumaniry 2r7-2r8, 234-240, social intelligence as preceding,

24r-243 r65; solar energy economy and,
superoxide dismutases, rz9 r99, 2oo, 2o2, 234-235; sp^ce
surplus, solar economy and, I9o-t9r, exploration, to-rz,r4, r3o, I3i,
r99-2O2, 2ro-2r2, 21 5, 234-235, r68, zr r, z16, 239-z4o; vision
242,246 and, 166
survival of the fittest, 19, 48, 53. See tectonic forces, zo6
a/so autopoiesis; extinction; natural tectonics, 206, pl. zj, pl. z6
selection teeth, r64
Swenson, Rod, r6-17 Teilhard de Chardin, Pierre, 5o, r7o
symbiogenesis: discovery of, xiii-xiv teleology. See purposefulness
Symbionticism and the Origin oJ Species, television, 186, r87, zz9
r33 telophase, pl. rz
symbiosis, pl. z, pl. t|a-b, pl. t9a; temperature maintenance, zz, 23, 32,
algaeand, Ioo, I20, t27,r33- 52. See also autopoiesis
134, r36, r77-t78, pl. tt, pl. t4; Tennyson, Alfred, 236
definition oi r r9; early work un- termites, tzo, 164, t68
covering, r 3 2- r 3 3 ; endosymbio- Tertiary period, r67
sis, defined, I I9-I20; food and, Tbtrahymena, zr8, zz4
r33-r34, r39-r4r, r44; individu- thallus, r74
ality and, tzo, 236-237;kchens thanosis (apoptosis), r37. See also
and, rzo, t77-t78; mitochondria aging; death
and,9o, I2r, I3o-r3r, r32-r33; thermodynamics: autopoiesis and, r9,
mitosis and, r23-r2g; molecular, 43, 8o-8r; laws of, r5-r7; and
75; of motiliry rz3, tz6-rzg; purposefulness, 16-17, 86, r5 r;
of organelles, t2r-r22; ts par- waste production and, r63
simonious explanation, rz8; thermophiles, 89, roo
plant-animal interdependence, Thermoplasma, too, 124, rz8-129,
2oo-2ot, 2o7-2ro: plastids and, r3o-r3r
r33-r3j; savanna evolution and, tholins, 74
zo8. See a/so autopoiesis; evolu- Thomas, Lewis, r r3
tion; protoctists tigers, rg3, r99,2oo
synergy, T, 8 Timaeus (Plato), 36
Szostak, Jack W., 84 timelines, 6z-6 3, 64-8o
tissue: diflerentiation of, r3J, I4o,
Thkami,Jun, 17r, r89 r j3-r 54, r jj; storage of energy
taxonomy. See classification of organic in, r9g
beings 'Irtan,74, to4
technology: agriculture, r35, r84- Tblypocladium, t88
r8J, zo8-2o9, 2ro, zrr; bacterial, toxics: animal recycling of, r63-t64;
238; and biosphere, t4,27, So, bacterial recycling of, I06-r08,
5 r-52, r68-t69, 216-218, 238- III
2 40, z 4r-2 43 ; biotechology, 93, Tiibrachidium, 16z
96, r8r, zo9; dependence on, Tiihonympha, 122, rz3
r86-187, 2to-ztr; evolution and, Tiichoplax, rj5, I56
zr6-2t8, 238-24o, z4t-243; and Tiidacna, r34
pollution, rc6, 164, r82,245- trilobites, r52, 160
246; purposefulness and, z3 r; truffles, t85-I86, r88
 288 lndex

truth: survival and, 3o-3 r wasps, r68

Tilber melanosporum, 18 s 'Wasson, R. Gordon, 17t, t87
Wasson, Valentina, r 87
ultraviolet radiation, ror; bacterio- waste. See pollution
phage production and, 96; ozone water: autopoietic planet and, zz-
layer and, ro6 z3; cyanobacteria and, ro3-r04;
uncertainty principle, 4 r-42 desert crusts and, zoz-zo3; ear\y
unconscious, 30, 22i-222, 223-224, Earth and, 6I; land-animal evolu-
226-229, 234 tion and, r67; oceans (see oceans);
undulipodia, tz4, rz6-127, tSS, r57, organisms as special form of, 5o;
166, zo3, pl. t7 origins of life ar7d,73,89; plant
United States, r87-r88, zoo evolution and. r95-r96. 2oz-2o3,
uraninite, ro5 zo4; purification of, zrr
lJranus, 6o water hyacinths, zt r
Urban VIII, rg Watson, James D., 7, 8
Urea, 24, z6 Watt, James, r 5
'Watts,
A1an,43
Varela, Francisco, r7 Way of All Flesh, The (B:utler), zz3
Veillard, S., r7z wealth, r9r, 2oo-2o2
Venus, zo, 2 1, 37, go Went, Frits, r95
Vernadsky, Vladimir Ivanovich, 48- whales, r68, r94
52, 55, r7o, rg3 What Is Lf? (Schrcidinger), r,7r
Viking mission, 12, r4 Whittaker, Robert H., r r9
viruses: autopoiesis vs., r8; bacterial Whittington, Harry B., r6o
reproduction and, 94, 95, 96; will, free, S, 2r7, 222, 23r-232.
RNA compared to, 84 See also choice
vision, ro3, t27, r65-167 will to live, 8o
vitalism, 3, 5, 7, 8, 17, 33, 4r, 2r7 Wilson, E. O., pl. 2 5 kdp.)
Vitousek, Peter, 2ro Woese, Carl,89, t3t
Vitreoscilla, g8 'Wohler, Friedrich, z4-25
Voltaire, 4o written language, r86-187, r89
Voluox, 136, pl. t4
Vostok I, to yeasts, rr6, ryz, 176, r88. See a/so
fungi
Walcott, Charles, r09, r6r
'Wallace, Alfred Russel,
45 Zea mays, 2o8-2og
'Wallin,
Ivan, r32-r33 zygomycotes, t76
'Walsh,
Maud, 88 zygotes, r40
Design Nola Burger
lllustrations Christie Lyons
Composition lntegrated Composition Systems
Text 11/14.5 Bembo
Display Syntax Regular and Bold
Printing and binding Friesens
lndex Victoria Baker
 SCIENCE / BIOLO6Y / CEOLOGY

Half a century ago, before the discovery of oNa, the Austrian "ln What ls Life? Margulis
physicist and philosopher Erwin Schrcidinger inspired a and Sagan have rephrased
generation of scientists by reframing the fascinating philo- the answer to Schrodinger's
sophical question: What is life? Using their expansive under- brilliant question by means of

standing of recent science to wonderful effect, acclaimed a new and spirited explanatio
of ,the emergent levels of
authors Lynn Margulis and Dorion Sagan revi5it this timeless
biological organization. . . .
question in a fast-moving, wide-ranging narrative that com-
Theirs is a conceptual frame-
bines rigorous science with philosophy, history and poetry.
work likely to influence futuri
The authors move deftly across a dazzling array of topics- introductions to biology."
from the dynamics of the bacterial realm, to the connecrion E. O. WILSON
befween sex and death, to theories of spirit and matter.
They delve into the origins of life, offering the startling sug- "A witty, exuberant panorama

gestion that life-not just human liG-is free to act and has of life that elaborates the plar
of symbiosis in evolution."
played an unexpectedly large part in its own evolution.
MARY CATHERINE BATESON
TLanscending the various formal concepts of life, this capti-
vating book offers a unique overview of lifet history, "This splendid book shows
essences, and future. how much more there is to lil
Supplementing the text are stunning illustrations that than mere reductionist biolog
range from the smallest known organism (Mycoplasrna bacte- Lynn Margulis and Dorion
ria) to the largest (the biosphere itself). Creatures both Sagan tread faithfully in Erwir
Schrodinger's footsteps and
strange and familiar enhance the pages of What Is Lfe? Their
are his true successors. "
existence prompts readers to reconsider preconceptions nor
JAMES E. LOVELOCK
only about life but also about their own part in it.
"A masterpiece of science
Lynn Margulis is Distinguished ProGssor in the Department writing. ... You will cherish
of Geosciences at the University of Massachusetts, Amherst, What ls llfe? because it is
and the recipient of the 1999 National Medal of Scien'e. so rich in poetry and science,
She is the author of more than one hundred articles and ten in the service of profound
books, including Symbiosis and Cell Euolution (second edition philosophical questions. "
1993). Dorion Sagan is the author of Biospheres (r99o). As MITCHELL THOMASHOW,

partners of Sciencewriters, Margulis and Sagan have also OR/ON

written Miuocosmos (California, 1996), What ls Sex? (rSqo),

Carden of Mkrobial Delights (rSqS), Mystery Dance (r99r), and
several guides to videos of live organisms.

ISBN 0-5-U-Ee0eI-8 LJnivcrsity of Cali&;rnia I)rcss

llc'rkeley 947:,o
llu llllil illilIlIl |]|| ttLt ll
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